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Full text of "Bulletin of the Natural Histort Museum. Geology series"

RD & W 2005 



ISSN 0968-0462 



Bulletin of 
The Natural Histor 
Museum 



HENA.tmrai 
j HISTORY MUSEUM 



Geology Series 




VOLUME 58 NUMBER 1 27 JUNE 2002 



The Bulletin of The Natural History Museum (formerly: Bulletin of the British Museum 
(Natural History) ), instituted in 1949, is issued in four scientific series, Botany, 
Entomology, Geology (incorporating Mineralogy) and Zoology. 

The Geology Series is edited in the Museum's Department of Palaeontology 
Keeper of Palaeontology: Dr N. MacLeod 

Editor of Bulletin: Dr M.K. Howarth 

Assistant Editor: Mr C. Jones 

Papers in the Bulletin are primarily the results of research carried out on the unique and ever-growing collections of the 
F m, both by the scientific staff and by specialists from elsewhere who make use of the Museum's resources. Many of the 

are works of reference that will remain indispensable for years to come. All papers submitted for publication are 
. :ed to external peer review for acceptance. 

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World list abbreviation: Bull. nat. Hist. Mus. Lond. (Geol.) 

Copyright © 2002 The Natural History Museum 

Geology Series Vol. 58, No. 1, pp. 1-79 
ISSN 0968-0462 

The Natural History Museum 

Cromwell Road 

London SW7 5BD Issued 27 June 2002 

Typeset by Ann Buchan (Typesetters). Middlesex 

Printed in Great Britain by Henry Ling Ltd, at the Dorset Press. Dorchester. Dorset 



Bull. not. Hist. Mus. Lond. (Geol.) 58( 1 ): 1-1 1 



Issued 27 June 2002 



Gough's Cave 1 (Somerset, England): a study 
of the axial skeleton 



■■ - ■■• ■-,■■ 



"CRY MUSEUM 

AUG 2002 



STEVEN E. CHURCHILL 

Department of Biological Anthropology and Anatomy, Duke University, Durham NC 27708, USA 

TRENTON W. HOLLIDAY 

Department of Anthropology, Tulane University, New Orleans LA 70118, USA 



r - 



PRESENTED 

PALAEONTOLOGY 



liBBARY 



SYNOPSIS. The postcranial axial skeleton of Cheddar Man (Gough's Cave 1 ) is represented by seventeen presacral vertebrae, 
the sacrum, and nineteen ribs, all of which are relatively well-preserved. Cheddar Man derives from early Holocene deposits in 
Gough's Cave, and the remains of his axial postcranial skeleton are described here. Comparative evaluation of the Gough's Cave 
1 remains reveals an axial skeleton that falls within the range of variation in size and shape of males of the same time period, albeit 
towards the small end of that range (reflecting relatively short stature in Cheddar Man). 



INTRODUCTION 



The postcranial axial skeleton of Cheddar Man is represented by a 
single cervical vertebra, eleven thoracic vertebrae, all five lumbar 
vertebrae, sacrum, and nineteen ribs. The hyoid, manubrium, ster- 
num and xiphoid process were not recovered. The preservation of the 
recovered vertebrae is generally good, with many being complete. 
Sequencing of vertebrae was based on size, details of morphology of 
the articular facets, neural arches, and transverse and spinous proc- 
esses, and by evaluating the articulation of each element with the 
identified supra- and subjacent vertebrae (Bass, 1987; Steele & 
Bramblett, 1988). Prior efforts to reconstruct the entire articulated 
vertebral column for museum display involved the gluing of fibrous 
pads (to represent the intervertebral discs) to the bodies of many of 
the vertebrae, and in some cases elements were glued to 'mocked up' 
replicas of the missing vertebrae, making observation and measure- 
ment of morphology difficult (only the fifth and eighth thoracic and 
the third lumbar vertebrae could be entirely separated from 
reconstructive materials: these specimens were thus singled out for 
photography). The ribs are also in a very good state of preservation 
overall; more than half of them preserve the head, neck and tubercles 
proximally and most are complete distally to the area around the 
anterior angle. Sequencing of ribs was accomplished by examining 
overall size and shape, the position of the M. iliocostalis line, size 
and shape of the articular facets, and the height of the rib heads (with 
the inferior bodies held in the same plane) relative to one another 
(Mann, 1993). A number of the ribs bear cut marks that may be 
attributable to stone tools. 

Each vertebra is briefly described, followed by a discussion of the 
vertebral morphology of Gough's Cave 1 (the sacrum is described 
along with the os coxae in Trinkaus, this series). The ribs are then 
likewise described and their morphology discussed. 



MATERIALS AND METHODS 



The description of the Gough's Cave 1 axial postcranial remains is 
augmented by osteometric data and comparisons with various sam- 
ples of fossil and recent humans. The necessity of accurate 
identification of vertebral and costal number (i.e., the position of the 
element in the series) for collection of comparative data presents 
difficulties in working with fragmentary fossil material (see 



Franciscus & Churchill, 2002). For vertebral morphology, compara- 
tive osteometries were collected on European terminal Pleistocene 
specimens (all associated with Late Upper Paleolithic assemblages, 
and dating between 19,000 and 11,000 ybp) and early Holocene 
specimens (associated with Mesolithic assemblages and dating 
between 10,000 and 5,000 ybp). These two samples thus bracket in 
age the Gough's Cave 1 skeleton. The terminal Pleistocene sample 
includes Arene Candide 2, 4, 5, 10 and 12, Bichon 1, Bruniquel 24, 
Cap Blanc 1 , Chancelade 1 , Grotta Contineza, Grotte des Enfants 3. 
La Madeleine, Oberkassel 1 and 2, Parabita 1 and 2, Le Peyrat 5 and 
6. Romito 4, St. Germain La Riviere 4 and Veyrier 1 (Paoli et al., 
1980; Simon & Morel, nd; Genet-Varcin & Miquel, 1967; von 
Bonin, 1935;Vallois, 1941^16, 1972;Verneau, 1906;deQuatrefages 
&Hamy, 1882; Verworn eta!., 1919; Cremonesi etal, 1972;Patte, 
1968; Graziosi, 1962; Vallois, 1972; Pittard & Sauter, 1945). The 
early Holocene sample is composed of Los Azules, Gramat 1, 
Hoedic8and9, Rastel LTeviec 1,11 and 16(Fernandez-Tresguerres, 
1976; Lacam et al, 1944; Barral & Primard, 1962; Pequart et al, 
1937). Additional comparative data was collected on recent Europe- 
ans (n = 125), north Africans (n = 61 ) and sub-Saharan Africans (n = 
26) (details of sample composition are provided in Holliday, 1995). 
For the ribs, comparative data is limited to a small sample of recent 
European-Americans (n = 20: Franciscus & Churchill, 2002). 

Operational definitions of the measurements employed can be 
found in Martin (1928) or as footnotes to Tables. Vertebral 
osteometries are provided in Tables 1-3, costal osteometries are in 
Tables 6-8. All measurements were taken by the authors on the 
original specimens; measurements quoted in brackets in Tables 1-9 
are estimated values. 



VERTEBRAL REMAINS 

Descriptions 

Cervical vertebra 6 or 7 (fig. 1 ) 

A single cervical vertebra, complete except for some damage to the 
left side ventral surface of the corpus, is preserved (at bottom of Fig. 
1 ). Based on its size and neural arch morphology (the transverse 
processes are large and laterally flaring) it appears to be either the 6th 
or 7th cervical vertebra (this element is attached superiorly to a 
'mocked up' cervical vertebral column, thus preventing examination 



) The Natural History Museum, 2002 




S.E. CHURCHILL AND T.W. HOLLIDAY 
Table 1 Dimensions (mm) of the sixth cervical vertebra. 



Dorso-ventral diameter 1 

Superior external transverse articular diameter 3 

Superior internal transverse articular diameter 1 

Superior transverse articular diameter 4 

Inferior external transverse articular diameter 5 

Inferior internal transverse articular diameter 6 

Inferior transverse articular diameter' 

Spinal canal dorso-ventral diameter (M-10) 

Spinal canal transverse diameter (M-l 1 ) 

Spinous process length 8 

Spinous process angle 1 * 

Body ventral height (M-l ) 

Body inferior dorso-ventral diameter (M-5) 

Body inferior transverse diameter (M-8) 



56.8 
51.7 

(31.5) 
41.6 
48.3 
21.8 
35.1 
14.7 
21.5 
29.2 
5° 

(12.2) 
17.3 
28.6 



Fig. 1 Gough's Cave 1 sixth or seventh cervical vertebra at bottom of 
figure, articulated to four reconstructed vertebra above; lateral view; x 1 . 



'From the mid- ventral surface of the body to the dorsal tip of the spinous process. 
2 Maximum distance between the lateral edges of the superior articular facets. 
'Maximum distance between the medial edges of the superior articular facets. 
""Average of the external and internal transverse articular diameters of the superior 
articular facets. 

'Maximum distance between the lateral edges of the inferior articular facets. 
6 Maximum distance between the medial edges of the inferior articular facets. 
'Average of the external and internal transverse articular diameters of the inferior 
articular facets. 

8 From the ventro-superior margin of the intersection of the laminae and the spinous 
process to the dorsal tip of the spinous process (not including the unfused tubercle). 
'The angle between the central long axis of the spinous process and the horizontal 
plane of the superior surface of the body, taken in the median sagittal plane of the 
vertebra. 

of the superior surface of the corpus and making its identification 
more difficult). The first thoracic vertebra is preserved, and it articu- 
lates poorly with this element, suggesting that this is the 6th cervical 
vertebra. The inferior surface of the body is concave (not flat as is 
normally found in 7th cervical vertebra: Bass, 1 987), and the anterior 
tubercle of the transverse process is relatively large and thus looks to 
be the carotid tubercle of C6. In addition, the end of the spinous 



Table 2 Dimensions (mm) of the thoracic vertebrae. 



Tl 



T2/3 



T4 



T5 



T6 



T7 



T8 



T9 T10 



Til 



ti: 



Dorso-ventral Diameter 1 

Superior external transverse articular diameter 

Superior internal transverse articular diameter' 

Superior transverse articular diameter 4 

Inferior external transverse articular diameter 5 

Inferior internal transverse articular diameter 6 

Inferior transverse articular diameter 7 

Spinal canal dorso-ventral diameter (M-10) 

Spinal canal transverse diameter (M-l 1 ) 

Spinous process length" 

Spinous process angle 1 * 

Body ventral height (M-l) 

Body dorsal height (M-2) 

Body median height (M-3) 

Body superior dorso-ventral diameter (M-4) 

Body superior transverse diameter (M-7) 10 

Body inferior dorso-ventral diameter (M-5) 

Body inferior transverse diameter (M-6) 10 



60.2 


- 


- 


- 


68.5" 


69.0" 


- 


- 


71.1" 


67.8" 


(69.2) 


46.1 


- 


36.2 


- 


31.4 


32.9 


33.1 


34.3 


35.9 


38.3 


- 


21.9 


- 


15.5 


15.2 


13.7 


- 


15.2 


14.7 


12.5 


15.1 


- 


34.0 


- 


25.9 


- 


22.6 


- 


24.2 


24.5 


24.2 


26.7 


- 


- 


38.0 


37.1 


33.9 


34.4 


34.5 


- 


37.5 


38.7 


- 


36.2 


- 


14.8 


13.8 


12.9 


- 


13.6 


- 


- 


10.4 


- 


17.3 


- 


26.4 


25.5 


23.4 


- 


24.1 


- 


- 


24.6 


- 


26.8 


16.7 


- 


16.5 


17.1 


15.2 


- 


17.8 


16.8 


- 


17.3 


- 


20.9 


(18.4) 


16.5 


17.1 


17.4 


17.1 


17.3 


18.2 


17.7 


20.0 


20.3 


31.3" 


(30)" 


- 


- 


36.4" 


37.8" 


- 


- 


36.3" 


29.1" 


26.3 


8° 


- 


- 


- 


47° 


70° 


- 


- 


55° 


45° 


16° 


13.7 


- 


17.3 


18.0 


17.0 


18.3 


19.4 


18.8 


19.1 


19.4 


(20) 


- 


- 


17.4 


18.1 


- 


- 


20.1 


- 


- 


- 


- 


- 


- 


- 


15.8 


- 


- 


19.4 


- 


- 


- 


- 


18.4 


- 


22.0 


23.2 


25.7 


- 


28.9 


32.8 


30.7 


32.2 


- 


30.2 


- 


26.7 


26.9 


29.7 


31.1 


32.2 


36.2 


36.5 


39.4 


- 


- 


- 


21.7 


25.6 


- 


30.2 


32.5 


- 


32.4 


- 


(34) 


- 


- 


28.5 


28.7 


31.3 


32.8 


36.4 


38.2 


39.3 


42.3 


41.7 



'From the mid- ventral surface of the body to the dorsal tip of the spinous process. 

-Maximum distance between the lateral edges of the superior articular facets. 

'Maximum distance between the medial edges of the superior articular facets. 

■"Average of the external and internal transverse articular diameters of the superior articular facets. 

'Maximum distance between the lateral edges of the inferior articular facets. 

"Maximum distance between the medial edges of the inferior articular facets. 

'Average of the external and internal transverse articular diameters of the inferior articular facets. 

"From the ventro-superior margin of the intersection of the laminae and the spinous process to the dorsal tip of the spinous process (not including the unfused tubercle). 

"The angle between the central long axis of the spinous process and the horizontal plane of the superior surface of the body, taken in the median sagittal plane of the vertebra. 

"Transverse body dimensions did not include the articular facets for the rib head. 

"Dorsal tubercle of spinous process unfused and missing. 



GOUGH'S CAVE AXIAL SKELETON 



process looks as though it gave rise to a bifid tubercle (the spinous 
process of the seventh cervical vertebra generally ends in a single 
tubercle (Williams & Warwick, 1980)), although the secondary 
centre of ossification is unfused and the process is preserved as a 
single tubercle. These features suggest that this bone represents the 
sixth cervical vertebra. 

The spinous process projects nearly horizontally from the body 
(Table 1), as is common in lower cervical vertebrae. The corpus is 
wide in the transverse dimension relative to its dorso- ventral diameter. 
As in all lower cervical vertebrae, the spinal canal is wider trans- 
versely than dorso- ventrally, and is triangular in outline. 

Thoracic vertebra 1 (fig. 2) 

The first thoracic vertebra is largely complete. The right side trans- 
verse process is broken off and the left side process is missing a small 
portion of its lateral end. The posterior tubercle of the spinous 





Fig. 3 Gough's Cave 
3c, inferior; x 1 . 



fifth thoracic vertebra. 3a, superior; 3b, lateral; 



Fig. 2 Gough's Cave 1 thoracic vertebrael-12 in articulation. 2a, lateral; 
2b, ventral; x 0.48. Note that reconstructed intervertebral disks of 
uniform thickness dorso-ventrally have been inserted between some of 
the vertebral bodies, probably diminishing the degree of curvature that 
would have obtained during life. 



process is unfused. A crack runs through the left side neural arch 
lamina and inferior and superior articular facets. The vertebra cannot 
be viewed from the inferior perspective due to adherent reconstructive 
materials. 

The spinous process projects nearly horizontally from the corpus 
(Table 2). The dorso- ventral diameter of the spinal canal is somewhat 
smaller than the transverse diameter. 

Thoracic vertebra 2 or 3 

The second or third thoracic vertebra is represented by the posterior 

portion of a neural arch only. This fragment includes the left side 



S.E. CHURCHILL AND T.W. HOLLIDAY 



lamina with the inferior articular facet, spinous process (with the 
posterior tubercle missing), and the right side lamina with the 
transverse process and superior and inferior articular facets. 

Thoracic vertebra 4 

This vertebra is complete except for the posterior half of the spinous 
process and the left side transverse process (Fig. 2). The superior and 
inferior surfaces of the body are obscured by reconstructive materials. 
The dorsal and ventral supero-inferior heights of the body are 
equal in this vertebra (Table 2). As with all of the thoracic vertebrae, 
the corpus has a greater transverse than dorso-ventral diameter 
(although the difference is not as great as that seen in the preserved 
[sixth or seventh] cervical vertebra). The dimensions of the spinal 
canal are equal in the transverse and dorso-ventral directions. 

Thoracic vertebra 5 (figs 2, 3) 

The fifth thoracic vertebra is complete except for most of the spinous 
process, the dorsolateral surface of the right transverse process, and 
the lateral end of the left transverse process (Fig. 3) 

As with the fourth thoracic vertebra, the dorsal and ventral supero- 
inferior heights of the body are equal (Table 2). The dimensions of 
the spinal canal are equal in the transverse and dorso-ventral direc- 
tions. 

Thoracic vertebra 6 

This vertebra is complete except for the lateral ends of both trans- 
verse processes (Fig. 2). This specimen is attached to the seventh 
thoracic vertebra inferiorly, and has reconstructive materials adher- 
ent to the superior surface of the body. 

The spinous process is infero-dorsally oriented (Table 2). The 
transverse diameter of the spinal canal is slightly greater than the 
dorso-ventral diameter in this element. 

Thoracic vertebra 7 

This element is complete except for the very tip of the spinous 
process and the lateral end of the right transverse process. This 
vertebra is affixed to the sixth thoracic vertebra superiorly and the 
inferior surface of the corpus is obscured by reconstructive material. 
The spinous process is strongly angled inferiorly (Table 2), and 
the transverse and dorso-ventral diameters of the body are sub-equal 
(with the transverse dimension being slightly larger). 

Thoracic vertebra 8 (figs 2, 4) 

This vertebra is mostly complete, lacking only the left side inferior 
costal facet (on the body), the right transverse process, the left 
inferior articular facet, and most of the spinous process (Fig. 4). The 
neural arch is cracked in several places and reconstructed. 

The dorsal supero-inferior height of the corpus is slightly greater 
than that of the ventral body (Table 2). The transverse and sagittal 
dimensions of the spinal canal are roughly equal in this vertebra. 

Thoracic vertebra 9 

The ninth thoracic vertebra is complete except for the end of the 
spinous process. The neural arch is broken off through both pedicles 
and has been reconstructed. This vertebra is attached to the tenth 
thoracic vertebra inferiorly and the superior surface of the corpus is 
covered by reconstructive material. 

The spinal canal of this vertebra shows an expansion of the 
transverse diameter and a diminution of the dorso-ventral diameter 
of the spinal canal relative to that of the suprajacent vertebra (Table 
2). 

Thoracic vertebra 10 

This vertebra is complete, but displays some slight damage to the 
right side inferolateral edge of the body. This specimen is attached to 
the ninth thoracic vertebra superiorly. 




Fig. 4 Gough's Cave 
4c, inferior; x 1 . 



eighth thoracic vertebra. 4a, superior; 4b, lateral; 



The spinous process is infero-dorsally directed, and is not as 
sharply inferiorly angled as that of the seventh thoracic vertebra 
(Table 2). 

Thoracic vertebra 1 1 

The eleventh thoracic vertebra is complete except for the very tip of 
the spinous process. The vertebra has some slight erosion to the 
inferior left side of the ventral surface of the corpus. The tip of the 
spinous process appears to be unfused. It is affixed to the twelfth 
thoracic vertebra inferiorly. 

The specimen exhibits slight anterior wedging of its body. The 



GOUGH'S CAVE AXIAL SKELETON 




Fig. 5 Gough's Cave 1 lumbar vertebrae in articulation. 5a, ventral; 5b, lateral; 5c, dorsal; x 0.72. Reconstructed intervertebral disks have been inserted 
between the lumbar bodies. 



spinous process has a more moderate inferior projection than that of 
the suprajacent vertebra (Table 2), and the spinal canal has a greater 
transverse than dorso- ventral diameter. 

Thoracic vertebra 12 

The twelfth thoracic vertebra is largely complete. This bone lacks 
only a portion of the left side ventral and lateral surfaces of the body, 
and the dorsolateral tip of the transverse process. The tip of the 
spinous process is unfused and missing, and the annular ring of the 
inferior surface is not fully fused to the centrum. The specimen is 
attached to the eleventh thoracic vertebra superiorly and its inferior 
surface is obscured by reconstructive material. 

The spinous process forms a moderate (inferiorly directed) angle 
with the plane of the body (Table 2). As with the lumbar vertebrae, 
the transverse diameter of the body is considerably greater than the 
dorso-ventral dimension. 

Lumbar vertebra 1 (fig. 5) 

The first lumbar vertebra is largely complete, lacking only the right 
side mammillary process. Some erosional damage is evident on the 
ventral surface of the body. The posterior tip of the spinous process 
appears to be unfused and missing (this region is obscured by 
reconstructive materials making observation of the morphology 



difficult). The bone is cracked through the right side pedicle and 
lamina and has been reconstructed. The specimen is attached 
inferiorly to the second lumbar vertebra. 

The body of the first lumbar vertebra exhibits marked anterior 
wedging (a much greater dorsoventral dimension inferiorly than 
superiorly: Table 3; Fig. 5). Erosion and damage to the anterior 
surface precludes measurement of the inferior dorsoventral diameter, 
and may accentuate the degree of wedging evident in the specimen. 
The spinous process is short and projects horizontally from the body 
(Table 3). The spinal canal transverse diameter is the largest of all the 
lumbar vertebrae, and is considerably greater than the dorso-ventral 
diameter. 

Lumbar vertebra 2 

This specimen is complete except for the lateral part of the right side 
transverse process (Fig. 5). The secondary centre of ossification for 
the tubercle of the spinous process is fused but the epiphyseal line is 
still open along its superior margin. The epiphyseal line between the 
secondary centre of ossification of the inferior annular ring and the 
centrum is also evident (but is mostly closed and was undergoing 
obliteration at the time of death). This bone is attached to the first 
lumbar vertebra superiorly and its inferior surface is covered by 
reconstructive material. 



S.E. CHURCHILL AND T.W. HOLLIDAY 



Table 3 Dimensions (mm) of the lumbar vertebrae. 



LI 



L2 



L3 



L4 



L5 



Dorso- ventral Diameter 1 

Superior external transverse articular diameter 

Superior internal transverse articular diameter' 

Superior transverse articular diameter 4 

Inferior external transverse articular diameter 5 

Inferior internal transverse articular diameter 6 

Inferior transverse articular diameter 7 

Spinal canal dorso-ventral diameter (M-10) 

Spinal canal transverse diameter (M-l 1 ) 

Spinous process length 8 

Spinous process angle 9 

Body ventral height (M-l) 

Body dorsal height (M-2) 

Body median height (M-3) 

Body superior dorso-ventral diameter (M-4) 

Body superior transverse diameter (M-7) 

Body inferior dorso-ventral diameter (M-5) 

Body inferior transverse diameter (M-6) 

Body sagittal angle 1 " 



80.2" 


85.1 


86.4 


- 


79.3 


35.7 


30.3 


34.8 


37.8 


54.7 


17.4 


- 


20.9 


20.3 


21.3 


26.6 


- 


27.9 


29.1 


38.0 


29.6 


31.0 


34.1 


46.2 


52.4 


- 


18.8 


16.4 


19.8 


27.9 


- 


24.9 


25.3 


33.0 


40.2 


18.9 


- 


15.2 


17.0 


18.3 


24.2 


20.9 


21.8 


22.6 


22.9 


27.8" 


33.9 


37.5 


- 


34.6 


2° 


2° 


16° 


- 


30° 


(20.3) 


22.0 


23.6 


26.2 


(25) 


(26.5) 


26.0 


26.9 


25.5 


24.0 


- 


- 


25.0 


- 


- 


(31.6) 


- 


40.3 


36.4 


34.1 


44.8 


45.5 


47.5 


50.0 


52.8 


- 


39.7 


37.2 


(36.4) 


(30) 


48.2 


50.3 


52.1 


52.9 


(50) 


-15° 


(-14°) 


-10° 


-6° 


+8° 



'From the mid-ventral surface of the body to the dorsal tip of the spinous process. 

2 Maximum distance between the lateral edges of the superior articular facets. 

'Maximum distance between the medial edges of the superior articular facets. 

•"Average of the external and internal transverse articular diameters of the superior articular facets. 

'Maximum distance between the lateral edges of the inferior articular facets. 

''Maximum distance between the medial edges of the inferior articular facets. 

'Average of the external and internal transverse articular diameters of the inferior articular facets. 

"From the ventro-superior margin of the intersection of the laminae and the spinous process to the dorsal tip of the spinous process (not including the unfused tubercle). 

"The angle between the central long axis of the spinous process and the horizontal plane of the superior surface of the body, taken in the median sagittal plane of the vertebra. 

"'Angle in the median sagittal plane between the tangents to the median sagittal surfaces of the superior and inferior vertebral disk surfaces (a positive angle has its apex 

dorsally and opens ventrally). 

"Dorsal tubercle of spinous process unfused and missing. 



The spinous process is of moderate length and is horizontally 
projecting from the corpus (Table 3). The dorsal supero-inferior 
body height is greater than that of the ventral surface, and the body 
is wide in transverse diameter relative to dorso-ventral diameter. 

Lumbar vertebra 3 (fig. 6) 

The third lumbar vertebra is complete except for the lateral portion 
of the right side transverse process. The tip of the spinous process is 
fused but the epiphyseal line is still open along its superior edge. The 
inferior and superior annular rings appear to be fully fused to the 
centrum, with the epiphyseal lines completely obliterated. 

The spinous process is mildly angled inferiorly relative to the 
plane of the corpus (Table 3) and is of moderate length. The body is 
supero-inferiorly higher on its dorsal than ventral aspect. Both the 
body and spinal canals are wide transversely relative to their dorso- 
ventral dimensions. 

Lumbar vertebra 4 

The fourth lumbar vertebra is largely complete. It lacks only the 
spinous process and the right side transverse process. Slight erosion 
to the ventral surface of the body is evident. The superior surface of 
the body is covered by reconstructive material. 

The ventral surface of the body is supero-inferiorly higher than the 
dorsal surface (Table 3). The corpus and spinal canal are transversely 
wide relative to their dorso-ventral diameters. 

Lumbar vertebra 5 

This vertebra is largely complete, lacking only the lateral ends of the 
transverse processes. Matrix is concreted to the left side transverse 
process, inferior articular facet and lamina. There is some erosion 
visible on the ventral surface of the body. The superior surface of the 
corpus is obscured by reconstructive material. 

The spinous process is shorter than that of the third lumbar 
vertebra (Table 3) and is the most inferiorly directed of all the lumbar 



vertebrae. The ventral surface of the body is higher supero-inferiorly 
than the dorsal surface. The body and spinal canal are transversely 
wide relative to their dorso-ventral diameters. 

Morphology 

When articulated, the thoracic vertebrae show a normal kyphosis 
(Fig. 2). The sum of the ventral body heights is 196.5 mm (using the 
average of the ventral heights of the first and third vertebrae for the 
missing corpus of vertebra 2), considerably shorter than the mean 
value for recent European males reported by Boule and Vallois 
(1937) of 243.1 mm, and is closer to the mean value of 221.9 mm 
obtained for European females (standard deviations and sample 
sizes not given). This is a reflection of the shorter stature of the 
Gough's Cave 1 individual relative to recent European males (see 
Holliday & Churchill, this series). The total ventral body height of 
Cheddar Man is more similar to, yet still on the small side of, the 
male and female skeletons from Teviec (male skeletons 2 [217.5 
mm] and 16 [231.0], female skeletons 1 [217.5] and 6 [223.5]: Boule 
& Vallois, 1937). This becomes more apparent when one looks at 
dorsal body heights, which are more reliable indicators of trunk 
height than are the ventral heights (which are frequently subject to 
anterior wedging: Stewart, 1966). Summary statistics for total tho- 
racic column height (summed dorsal body heights for T1-T12) for 
comparative samples can be found in Table 4. The total thoracic 
height figure for Gough's Cave 1 and the majority of the fossil 
sample were predicted via least-squares regressions of total thoracic 
height on those elements preserved for a recent human series (n=45: 
Holliday, 1995). The standard error of the estimate for the measure- 
ments predicted by this method is very low (Holliday, 1995), 
providing a reasonable degree of confidence in the predicted values. 
In no case was a predicted thoracic height used if its standard error of 
the estimate was greater than 3% of the prediction itself. 



GOUGH'S CAVE AXIAL SKELETON 




Table 4 Summary statistics for thoracic column height (mm) in Recent 
and Late Pleistocene/Early Holocene samples (mean; SD; n). 



Fig. 6 Gough's Cave 1 third lumbar vertebra. 6a, superior; 6b, lateral; 6c, 
inferior; x 0.9. 

As is evident from Table 4, Gough's Cave 1 has a short thoracic 
column relative to most of the mean values for males in the compara- 
tive samples. The value for his thoracic height is less than that of all 
the male means, with the exception of the recent sub-Saharan 
Africans. His value falls within the low end of the male range for 
most of the samples. Interestingly, his thoracic height falls very close 
to the mean values of all of the European (Pleistocene, Holocene and 
recent) female samples. This of course reflects the overall short 
stature of the Gough's Cave 1 specimen. 





Total Thoracic Height 




Male 


Female 


Gough's Cave 1 


234.7 




Mesolithic Europeans 


25 1.7; 20.4; 4 


233.8; 5.7; 2 


Late Upper Paleolithic Europeans 


256.8; 18.1; 12 


231.8; 10.1:3 


Recent Europeans 


259.0; 12.9; 63 


238.3; 12.6; 52 


Recent North Africans 


239.9; 15.8; 26 


223.5; 8.4; 28 


Recent Sub-Saharan Africans 


229.6; 11.0; 9 


219.7; 16.2; 15 



When articulated, the lumbar vertebrae evince a normal lordosis 
(Fig. 5). The sum of the ventral body heights is more similar to those 
observed in the males from Teviec with five lumbar vertebrae (118.5 
mm in Teviec 2, 1 16.0 in Teviec 4). The females from Teviec with 
five lumbar vertebrae have total ventral body heights that are slightly, 
but not substantially, shorter (1 10.0 mm in Teviec 1, 1 12.0 mm in 
Teviec 3). Three of the skeletons from Teviec have six lumbar 
vertebrae, without a reduction in the number of thoracic vertebrae, 
and thus have lumbar regions that are substantially longer supero- 
inferiorly (Teviec 16 [male], 155 mm; Teviec 6 [female] 148.5 mm: 
Boule & Vallois, 1937). As with the thoracic vertebral column, 
inclusion of the summed dorsal body heights of the lumbar vertebrae 
allows the comparison of Gough's Cave 1 to several Recent and 
fossil human samples. As for the thoracic column heights, lumbar 
column heights were predicted via least-squares regressions; none 
was used if its standard error of the estimate exceeded 3% of the 
predicted measurement. Table 5 shows that Cheddar Man has a 
shorter lumbar column than the male mean of all but one compara- 
tive sample (recent North Africans). To some extent, this is due to the 
marked posterior wedging exhibited in the specimen's L3-L5 verte- 
brae (see below). However, his relatively small size also plays a role; 
his lumbar column height falls squarely among the means for all the 
European female samples. Importantly, the male mean for the 
Mesolithic sample is high due to the inclusion of Teviec 16, who, as 
discussed above, has 6 lumbar vertebrae. 

In recent Europeans, the ventral body height is typically greater 
than the dorsal body height in the fourth and fifth, and often in the 
third, lumbar vertebrae (Boule & Vallois, 1937). In the sample from 
Teviec, this pattern generally holds only for the fifth vertebra (Boule 
& Vallois, 1937). Gough's Cave 1 evinces the pattern seen in recent 
Europeans, with a greater supero-inferior dimension of the ventral 
body in the third, fourth and fifth lumbar vertebrae (Table 3). The 
lumbo-vertebral index (100 * [I dorsal body heights]/[E ventral 
body heights]) is 110.1 in Cheddar Man, higher than the mean value 
for the Teviec specimens but not outside their range (mean of six 
individuals = 103.6, range 96.3-110.1: Boule and Vallois, 1937). 
The position of the articular facets of the vertebrae indicate a lordotic 
curvature to the lumbar column (with perhaps greater lordosis cre- 
ated in the lower lumbars: Fig. 5), so there must have been 
considerable wedging of the intervening intervertebral disks. 

Table 5 Summary statistics for lumbar column height (mm) in Recent 
and Late Pleistocene/Early Holocene samples (mean; SD; n). 





Total Lumbar Height 




Male 


Female 


Gough's Cave 1 


128.9 




Mesolithic Europeans 


137.4; 13.5:4 


126.9:8.4:2 


Late Upper Paleolithic Europeans 


130.1; 8.6; 11 


127.9; 2.5; 4 


Recent Europeans 


134.9; 8.0; 66 


128.2:7.4:59 


Recent North Africans 


127.0; 10.5:29 


123.4:7.3:32 


Recent Sub-Saharan Africans 


131.5:5.9: 11 


122.7:9.0; 15 



S.E. CHURCHILL AND T.W. HOLLIDAY 



The spinous processes of the thoracic and lumbar vertebrae are 
unremarkable and not particularly robust, similar to the condition 
observed in the Teviec skeletons (Boule and Vallois, 1937). The 
transverse processes of the lower thoracic vertebrae are, however, 
relatively large and robust. The insertion areas for the levator costae 
muscles and costotransverse ligaments tend to be well marked on the 
ribs (see below), suggesting some overall robusticity in the thorax of 
Cheddar Man (at least with respect to muscles and structures import- 
ant in respiration). The inferior demi-facets for the rib heads on the 
centra are quite large in most of the thoracic vertebrae, and tend to 
form laterally-projecting tubercles with inferiorly directed articular 
surfaces. The flattening of the left side ventral bodies that usually 
occurs in thoracic vertebrae 5-8 (from pressure from the aorta) is 
only slightly apparent in Gough's Cave 1 . 



COSTAL REMAINS (FIG. 7) 

Descriptions 

Rib 2 

The right second rib is preserved as a 78.8 mm-long fragment from 

the neck just proximal of the tubercle to the region of the proximal 



end of the M. serratus anterior tubercle (the proximal part of the 
tubercle is apparent). The left second rib is preserved as a 95.9 mm- 
long fragment from mid-neck to just distal of the M. serratus 
anterior tubercle, and the superior surface of the distal half of the 
fragment is covered with a thin layer of matrix (Fig. 7). 

The right-side rib has a well developed crest for M. scalenus 
posterior and a distinct groove on the external edge of the inferior 
surface for the intercostal muscles and membranes. The M. scale- 
nus posterior crest is not as strongly developed on the left-side rib 
(although the difference is slight), but the region just internal of 
the crest (on the superior surface) is more rugose. The M. serratus 
anterior crest on the left rib is very weakly developed. A piece of 
the superior surface of the shaft of this rib is missing in the region 
of the proximal tubercle, and the rest of the tubercle is covered by 
thin matrix, but it is clear nonetheless that the tubercle is not 
large. The left rib also displays a distinct groove on the external 
edge of the inferior surface for the intercostal muscles and 
membranes, but it is not as well defined as on the right side rib. 
The non-articular tubercles are relatively slight, with the one on 
the left rib being slightly larger. The articular facets (measuring 
10.2 mm proximodistally (PD) by 6.1 mm supero-inferiorly (SI) 
on the right and 9.1 PD by 6.3 SI on the left) are dorsoinferiorly 
directed. 




Fig. 7 Gough's Cave 
of the photograph. 



ribs in superior view; x 0.43. The ribs are arranged in sequential order with the second ribs at the top and right-side ribs to the right 



GOUGH'S CAVE AXIAL SKELETON 
Table 6 Dimensions (mm) of ribs 2-4. 



R2 



L2 



R3 



L3 



R4 



Neck length' 
Proximal thickness (M- 
Proximal height (M-l) 
Shaft thickness 3 
Shaft height 1 



11.2 
7.3 



12.2 
7.1 



(11.7) 4 (12.3) 4 



8.4 
8.3 



(27.9)- 
8.9 
9.3 



7.7 



(7.1) 4 (7.3) 4 11.7 10.1 7.7 



'Distance from the middle of the head to the middle of the articular tubercle. 
2 Head unfused, measurement taken from middle of epiphyseal surface for head. 
'Rib thickness (internal-external diameter, measured in the plane of rib curvature) 
and height (supero-inferior diameter, taken perpendicular to the plane of curvature of 
the rib) at the point where the M. iliocostalis line meets the inferior edge of the rib. 
4 Taken 1 cm distal of where proximal thickness and height were taken. 

RIB 3 

The right-side third rib is represented by a 154.7 mm-long fragment 
from mid-neck to just proximal of the anterior angle. The left third 
rib is preserved as a 171 .7 mm-long fragment from the distal end of 
the posterior angle to just proximal of the sternal end. 

The M. iliocostalis line 1 in the right rib is not pronounced (a 
feature of all of the Cheddar Man's ribs). A small portion of the M. 
iliocostalis line is preserved proximally in the left-side rib, and it 
looks to have been more strongly developed than in the right (how- 
ever, the rib itself is somewhat slighter). The right rib shows a 
discernable attachment for M. levator costae and both ribs have a 
distinct sulcus on the superior edge of the rib in the vicinity of the 
posterior angle (ca. 30 mm long) for the intercostal muscles. There is 
no discernable subcostal groove on the right rib, and the left side 
shows a weak subcostal groove for only a few centimeters distal of 
the posterior angle. The left rib has a supero-inferior flare to the body 
about 45 mm proximal of the anterior angle, reflecting perhaps a 
healed fracture. The articular facet on the right side is dorsoinferiorly 
directed and measures 9.5 mm (proximodistally) by 7.8 mm (supero- 
inferiorly). 

Rib 4 

The right fourth rib is a 1 27.0 mm-long fragment preserved from the 
head to somewhere proximal of midshaft. The proximal end of the 
rib is intact. The left fourth rib is a 1 56.8 mm-long fragment of the rib 
body, from somewhere distal of the posterior angle to the region of 
the anterior angle. 

On the right side, the surface of the head is rough and irregular, 
likely representing the subchondral surface of the unfused secondary 
centre of ossification for the head. There is a small and superiorly 
directed tubercle on the neck, and from this a crest runs distally along 
the superior margin of the bone past the non-articular tubercle, most 
likely representing the attachment of the superior costotransverse 
ligament. The M. iliocostalis line is not pronounced. The articular 
tubercle is large (9.0 mm proximodistally by 11.5 mm internal- 
external) and is primarily inferiorly directed. The subcostal groove is 
weakly developed on both ribs. In the right rib there is a strong bend 
at the posterior angle (the angle between the head-neck axis and the 
proximal costal body is approximately 90°). 

Rm5 

The right fifth rib is preserved as a 190 mm-long fragment, intact 
from the head down to the anterior angle, and missing only a portion 
of the sternal end. The left rib is represented by a 210 mm-long 



fragment, also intact from the head down to the anterior angle and 
missing only a part of the sternal end. 

The secondary centres of ossification for the heads are only 
partially fused (and portions are missing) on both sides. As in the 
right fourth rib. the fifth ribs present small superiorly directed 
tubercles on the neck that continue distally as crests running along 

Table 7 Dimensions (mm) of ribs 5-7. 



R5 



L5 



R6 



L6 



R7 



L7 



Rib length (M-4) - >200' 

External arc (M-3) - >323' - 

Neck length 2 (27.3) 3 (28.5)-' (23.9)' (25.3) 1 - (26.9) 5 

Proximal thickness (M-2) 9.2 8.3 8.0 8.6 - 8.9 

Proximal height (M-l) 8.6 8.7 10.0 12.0 - 9.3 

Shaft thickness 4 9.3 9.6 8.6 8.7 9.7 9.3 

Shaft height 4 13.0 14.5 14.4 13.6 13.5 

Chord 5 - (213)' 

Subtense 6 (77)' 

Transverse width 7 8.2 

'Rib is missing a small portion of the sternal end. 

^Distance from the middle of the head to the middle of the articular tubercle. 

'Head unfused, measurement taken from middle of epiphyseal surface for head. 

J Rib thickness (internal-external diameter, measured in the plane of rib curvature) 

and height (supero-inferior diameter, taken perpendicular to the plane of curvature of 

the rib) at the point where the M. iliocostalis line meets the inferior edge of the rib. 

'Distance from the distal margin of the articular tubercle to the proximal extent of 

the sternal end, following McCown and Keith (1939: fig. 75). 

''Maximum perpendicular distance from the chord to the external surface of the rib. 

following McCown and Keith (1939; fig. 75). 

'Internal-external diameter of the rib body at the intersection of the subtense. 



the superior margins of the bones past the non-articular tubercles. 
These crests, most likely marking the sites of attachment of the 
superior costotransverse ligaments, are more strongly developed 
than that on the fourth rib. Both ribs also have a crest on the inferior 
edge of the neck running from the head to the proximoinferior edge 
of the articular facet. These crests may represent the attachments of 
expanded accessory ligaments from the heads and necks of the 
subjacent ribs (Williams & Warwick, 1980) or distal extensions of 
the radiate ligaments binding the heads to the adjacent vertebra. The 
nonarticular tubercles are bulbous and projecting, and the articular 
tubercles are inferodorsally directed (measuring 7.9 mm PD by 10.3 
mm SI on the right, and 7.2 mm PD by 10.5 mm SI on the left). The 
M. iliocostalis lines are not very well developed and it is hard to make 
out where the lines cross the inferior border of the rib. The subcostal 
grooves are neither deep nor strongly developed but are clearly 
visible along most of the body. The angle between the head/neck axis 
and the axis of the body is about 1 17° on both sides. 

Rib 6 

The right sixth rib is preserved as a 165.3 mm-long fragment, 
complete from the head to somewhere distal of midshaft. The left- 
side rib is a 1 85. 1 mm-long fragment, complete from the head to the 
area of the anterior angle. 

The centres of ossification of the heads are incompletely fused and 
portions of them are missing. The ribs of both sides have short necks 
with small tubercles on their superior surfaces for the superior 
costotransverse ligaments. The ribs lack the crests (distal of the 
tubercles) that are seen on the suprajacent ribs. The inferior edges of 



'The iliocostal muscles are the lateral-most extensions of the erector spinae {sacrospinal!: ) muscle. The M. iliocostalis lumborum inserts on the inferior borders of the lower six or 
seven ribs, at the posterior angle (Williams & Warwick, 1980). M. iliocostalis thoracis arises from the superior borders of the angles of the lower six ribs and inserts on the superior 
margins of the upper six ribs. M. iliocostalis cervicis attaches to the superior borders of the third to sixth ribs. Thus various combinations of these muscles, as well as the thoracolumbar 
fascia, contribute to the formation of the iliocostalis lines on the external surface of the posterior angle of the ribs, and will be referred to throughout this description as the iliocostalis 
muscle. 



10 



S.E. CHURCHILL AND T.W. HOLLIDAY 



the necks also lack the crests seen on the fifth ribs. The articular 
tubercles are primarily interiorly directed and round in shape (right- 
side 9.7 mm PD by 9.5 mm internal-external (IE) , left-side 9.4 mm 
PD by 9.5 mm IE). The nonarticular tubercles are almost absent, 
appearing as small dorsosuperior extensions of the articular facets. 
The M. iliocostalis lines are not rugose nor marked. The attachments 
for M. levator costae can be seen as crests on the superior edges of 
the ribs running distally from the level of the tubercles and blending 
into the M. iliocostalis lines. The subcostal grooves are clear and 
distinct. The head/neck axis to shaft axis angle is roughly 135° in 
both ribs. 

R1B7 

The right seventh rib is preserved as a 181.1 mm-long fragment of 
the body, from somewhere distal of the posterior angle to just distal 
of the anterior angle. Judging from the size and curvature of the left 
side antimere, the proximal break occurred right at the distal end of 
the posterior angle. The left-side rib is represented by a 185.0 mm 
long-fragment, complete from the head to the region of the anterior 
angle. 

The secondary centre of ossification for the head of the left rib is 
unfused and missing. The neck is short and has a tubercle and crest 
on its superior surface for the superior costotransverse ligament. The 
neck has a large foramen or pit (plugged with matrix) on its dorsal 
surface. The M. levator costae crest is pronounced. The articular 
facet is dorsoinferiorly directed and oval in shape (10.8 mm PD by 
8.4 mm IE). The nonarticular tubercle is poorly defined but is larger 
than that of the 6th rib. The M. iliocostalis line is non-rugose and 
poorly defined. On both sides the proximal bodies are quite thick and 
form a 'roof over the subcostal groove along the proximal shaft. The 
subcostal grooves are very clearly defined along the proximal por- 
tions of the shafts. In the left-side rib, the head/neck axis to body axis 
angle is strong (approximately 109°). 

Rib 8 

The right eighth rib is preserved as a 198 mm-long fragment, 
complete from the head to the area of the anterior angle. The left rib 
is preserved as a 190.3-mm long fragment of the body, retaining a 
small portion of the nonarticular tubercle proximally. Distally the 
left-side rib is broken somewhere proximal of the anterior angle. 

On the right rib the head is unfused and missing. The neck is short 
and has a tubercle and crest on its superior surface for the superior 
costotransverse ligament. This crest is continuous with the insertion 
of M. levator costae and blends with the superior portion of the M. 
iliocostalis line distally. The same morphology can be seen in the 
left-side rib from the area of the M. levator costae insertion (the most 
proximally preserved portion of the shaft) down to the M. iliocostalis 
line. The inferior surface of the right-side neck also has a clearly 
defined crest, perhaps reflecting the attachment of an expanded 
accessory ligament from the head and neck of the subjacent rib 
(Williams & Warwick, 1980) or a distal extension of the radiate 

Table 8 Dimensions (mm) of ribs 8-11. 





R8 


L8 


R9 


L9 


Rll 


Lll 


Neck length 1 


(26.7) 2 


_ 


(22.5) 3 


(23.1)- 


_ 


_ 


Proximal thickness (M-2) 


10.2 


9.3 


8.4 


8.2 


- 


- 


Proximal height (M-l) 


9.8 


9.2 


9.7 


9.1 


- 


- 


Shaft thickness 3 


- 


10.6 


- 


7.9 


6.9 


6.3 


Shaft height 3 


- 


11.4 


- 


15.3 


13.2 


13.8 



'Distance from the middle of the head to the middle of the articular tubercle. 
-Head unfused. measurement taken from middle of epiphyseal surface for head. 
'Rib thickness (internal-external diameter, measured in the plane of rib curvature) 
and height (supero-inferior diameter, taken perpendicular to the plane of curvature of 
the rib) at the point where the M, iliocostalis line meets the inferior edge of the rib. 



ligament. The neck has a foramen or pit on the dorsal surface just 
proximal to the articular facet. The articular facet is oval (11.6 mm 
PD by 9.3 mm IE) and is primarily inferiorly directed. The 
nonarticular tubercle is poorly defined and blends with the proximal 
end of the M. iliocostalis line. The M. iliocostalis lines are non- 
rugose and poorly defined on both ribs. Small, mildly rugose 
depressed areas can be seen on the superior margins of the shafts just 
distal to the M. iliocostalis lines, likely marking the proximal extent 
of the insertion of the intercostal muscles. The subcostal grooves are 
very clear along the proximal halves of the ribs. The head/neck axis 
to body axis angle of the right-side rib is 122°. 

Rib 9 

The right ninth rib is a 79.6 mm-long fragment of the proximal end, 
complete from the head to the region of the posterior angle. The left 
rib is preserved as a 150.5 mm-long fragment, complete from the 
head to somewhere below midshaft. 

The centres of ossification for the heads are unfused and missing. 
The necks are relatively short. In the right-side rib, a small tubercle 
is evident on the superior surface of the neck, perhaps reflecting an 
attachment for an accessory ligament that ran superiorly to the crest 
on the inferior surface of the neck of the right eighth rib. The crests 
for M. levator costae are clearly defined on both ribs. The articular 
facets are oval shaped (measuring 8.9 mm PD by 7.7 mm IE on the 
right, 8.9 mm PD by 8.0 mm IE on the left) and are inferodorsally 
directed. The nonarticular tubercles arise from the articular tubercles 
and are relatively small. The M. iliocostalis lines are not pronounced. 
The costal grooves are wide supero-inferiorly and shallow. The 
head/neck axis to shaft axis angle is 116° in both ribs. 

Rib 11 

The right eleventh rib is represented by a 130.5 mm-long fragment, 
complete from the head to somewhere near the anterior angle. The 
left-side rib is preserved as a 102.5 mm-long fragment, complete 
from the head to 37 mm below the posterior angle. The shaft of the 
left rib is eroded and damaged. 

The heads are unfused and missing on both sides. The ribs present 
neither articular nor nonarticular tubercles, but crests for the cos- 
totransverse ligaments are visible on the superior margins of the 
proximal bodies. Narrow, oval insertion scars for the intercostal 
muscles are visible on both the inferior and superior edges at the 
posterior angle. The M. iliocostalis lines are indistinct, but bulging 
tubercles are visible at the posterior angle in each rib for the attach- 
ment of this muscle. 

Rib 12 

The right twelfth rib is complete, and has a total length of 96.6 mm. 
The area of the internal intercostal muscle attachment on the superior 
edge is eroded away, but otherwise the bone is well preserved. 

The head appears to be unfused. The rib has a clear crest on the 
superior surface of the neck for the costotransverse ligament. There 
is also a sulcus on the inferior edge of the internal surface of the 
proximal shaft for M. quadratus lumborum. The diaphragm attach- 
ment is indistinct. There is a long (19 mm), narrow scar on the 
inferior edge of the distal shaft forM serratus posterior inferior, and 
on the superior external surface some rugosity is visible that may 
represent the attachment site of M. latissimus dorsi. The right twelfth 
rib is 95.9 mm long (M-4: Martin, 1928) and has an external arc 
length (M-3: Martin, 1928) of 54 mm. 

Unidentified shaft fragment 

This is a 58.2 mm-long fragment of the distal end of a rib body, 
including the sternal end. The subcostal groove is not preserved, and 
thus the side cannot be determined. The fragment is somewhat 
damaged and has some areas of plaster reconstruction. The supero- 



GOUGH'S CAVE AXIAL SKELETON 



11 



inferior height of the body is 14.4 mm. Based on the size and shape 
of the fragment, it appears to be part of an upper rib. 

Morphology 

The overall size (Tables 6-8), shape, robusticity and muscularity of 
the Gough's Cave 1 ribs fall within the range of variation of recent 
human samples. In terms of rib shaft height and thickness the 
Gough's Cave 1 ribs generally fall within one standard deviation of 
the mean values obtained in the comparative sample (Table 9). Rib 
shaft shape, as measured by the ratio of thickness to height, is also 
generally within one standard deviation of the Euro American means. 
The notable exceptions concern the fourth and eighth ribs, both of 
which are markedly shorter in the supero-inferior dimension, result- 
ing in shaft shape ratios that are elevated (indicating a 'rounder' shaft 
cross-section) relative to the comparative sample. 

The M. iliocostalis lines are generally poorly marked in the entire 
series of ribs. Other muscle markings and ligamentous insertions 
tend to be more pronounced. Most of the ribs exhibit a distinct crest 
for M. levator costae and clear attachment areas for the superior 
costotransverse ligament. However, the insertions on the external 
surfaces of the rib necks for the costotransverse ligaments are not 
pronounced, and, with the exception of the fifth rib, the non-articular 
tubercles (upon which the lateral costotransverse ligaments attach) 
are slight. Evidence of expanded accessory or radiate ligaments can 
be seen in several of the ribs. The subcostal grooves tend to be 
weakly developed in the upper ribs of the series, but are distinct in the 
middle ribs. The insertion sites of the intercostal muscles are well 
defined in a number of the ribs. The insertion of M. scalenus 
posterior is evident in both second ribs, yet the M. serratus anterior 

Table 9 Rib shaft dimensions (mm) in Gough's Cave 1 and recent 
European-American males (mean, SD). 







EuroAmerican Males 


RIB 


Gough's Cave 1 


(n = 20) 


2 Thickness 


(11.7)' 


12.7 ± 1.1 


Height 


(7.1) 1 


7.3 ±0.8 


T/H ratio 


1.65 


1.73 ±0.2 


3 Thickness 


8.3 


7.8 ±1.1 


Height 


11.7 


11.2± 1.7 


T/H ratio 


0.71 


0.71 ±0.1 


4 Thickness 


8.1 


8.6 ±0.9 


Height 


7.7 


11.7 ± 1.9 


T/H ratio 


1.05 


0.74 ±0.1 


5 Thickness 


9.3 


9.0 ± 1.0 


Height 


13.0 


12.8 ±1.6 


T/H ratio 


0.72 


0.71 ±0.1 


6 Thickness 


8.6 


9.2 ± 1.0 


Height 


14.5 


13.9 ± 1.5 


T/H ratio 


0.59 


0.67 ±0.1 


7 Thickness 


9.7 


9.0 ±1.0 


Height 


13.6 


15.0± 1.9 


T/H ratio 


0.71 


0.61 ±0.1 


8 2 Thickness 


10.6 


8.6 + 0.8 


Height 


11.4 


15.5 ±2.5 


T/H ratio 


0.93 


0.57 ±0.1 


9 2 Thickness 


7.9 


8.0 ±0.8 


Height 


15.3 


17.0±2.8 


T/H ratio 


0.52 


0.48 ±0.1 


1 1 Thickness 


6.9 


6.1 ± 1.0 


Height 


13.2 


12.9 ±1.6 


T/H ratio 


0.52 


0.48 ±0.1 



'Taken 1cm distal of location of proximal thickness and height measurements. 
! Taken on left-side rib for Gough's Cave 1. 



tubercle is relatively slight in the left side rib (this region is not 
preserved in the right-side rib). In the preserved right twelfth rib, an 
obvious sulcus for M. quadratus lumborum can be seen, and the 
attachment areas of M. serratus posterior inferior and M. latissimus 
dorsi are clearly evident. The presence of crests for the levator costae 
muscles, along with the visible insertion sites of the intercostal 
muscles in some ribs and the development of the attachment areas of 
the ligaments that bind the ribs to vertebrae suggest a moderately 
high level of respiratory activity in this individual. However, this 
general robusticity does not extend to all of the muscles of the back 
and trunk that arise from or attach to the ribs. 



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Bull. nat. Hist. Mus. Lond. (Geol.) 58(1): 13-79 Issued 27 June 2002 

Upper Ordovician brachiopods from the 
Anderken Formation, Kazakhstan: their 
ecology and systematics 

L.E. POPOV 

Department of Geology, National Museum of Wales, Cardiff CF 10 3NP 

L.R.M. COCKS 

Department of Palaeontology, The Natural History Museum, Cromwell Road, London SW7 5BD 

I.F. NIKITIN 

Institute of Geological Sciences, Almaty 480100, Kazakhstan 

CONTENTS 

Synopsis 13 

Introduction 13 

Outline of geology and fossil localities 14 

Faunal Associations 22 

Overall palaeoecology 26 

Systematic palaeontology 27 

Linguloidea 28 

Discinoidea 28 

Siphonotretoidea 30 

Craniopsoidea 30 

Strophomenoidea 30 

Plectambonitoidea 38 

Chilidiopsoidea 53 

Triplesioidea 53 

Protorthoidea 58 

Orthoidea 58 

Plectorthoidea 61 

Enteletoidea 64 

Camarelloidea 64 

Rhynchotrematoidea 74 

Lissatrypoidea 76 

Meristelloidea 76 

Acknowledgments 77 

References 77 

SYNOPSIS. The brachiopod fauna from the Anderken Formation ( Lower to Middle Caradoc) of the Chu-Ili Range, south-eastern 
Kazakhstan, is revised and described systematically. It consists of 62 species in 55 genera, of which the genera Tesikella, 
Olgambonites and Zhilgyzambonites (all Plectambonitoidea) and llistrophina (Camarelloidea) are new, and the species Bellimurina 
(Bellimurina) sarytumensis, Teratelasmella chugaevae, Foliomena prisca, Acculina kulanketpesica, Dulankarella larga, Kajnaria 
rugosa, Anoptambonites convexus, Olgambonites insolita, Zhilgyzambonites extenuata, Gacella institata, Placotriplesia spissa, 
Grammoplecia wrighti, Dolerorthis pristiua, Austinella sarybulakensis, Plectorthis ? burultasica, Bowanorthis? devexa, Pionodema 
opima, Parastrophina iliana, llistrophina tesikensis, Liostrophia pravula, Plectosyntrophia unicostata, Rhynchotrema akchokense 
and Nikolaispira guttula are new. Six brachiopod-dominated assemblages are recognised and defined, termed the Ectenoglossa, 
Tesikella, Mabe/Ia-Sowerbyella, Acculina-Dulankarella, Parastrophina- Kellerella and Zhilgyzambonites-Foliomena Associa- 
tions. The relationships with contemporary faunas are assessed, and the Anderken brachiopods appear to have much in common 
with those of north-west China. 



INTRODUCTION 



_ _ separate crustal fragments of variable size. The relative positions of 

these fragments are contentious; some authors, notably Sengor & 

Natalin (1996), consider that most of the fragments made up an 

The global geography of the Lower Palaeozoic has been the subject enormous island arc. termed the Kipchak Arc, which stretched in a 

of widespread international discussion in recent years (references in long curve all the way from the substantial craton of Baltica to the 

Cocks 2001 ), but much is not yet clear. In the Ordovician, the large central Siberian Angaran craton. Others, for example Nikitin, have 

area of what is today Kazakhstan was then divided into many subdivided Kazakhstan in other ways, with a more conservative 

© The Natural History Museum, 2002 



14 



L.E. POPOV, L.R.M. COCKS AND I.F. NIKITIN 




Fig. 1 Generalised map of the Chu-Ili Range and West Balkhash Region (including the southern part of Lake Balkhash), showing the boundaries of the 
Early Palaeozoic tectonofacies belts, mainly after Nikitin et al. ( 1991 ), and the position of the brachiopod localities discussed in the text: 1, Anderkenyn- 
Akchoku; 2, Kujandysai; 3, east side of Kopalysai River; 4, Buldubai-Akchoku Mountain; 5, Tesik River; 6, Burultas Valley; 7, south-east side of Karatal 
River near Sorbulak spring; 8, 7 km southwest of Karpkuduk Well, Kotnak Mountains. 



palaeogeography. Up until now, little assessment of the faunas 
contained within these tectonic plates has been made, particularly in 
relationship to contemporary faunas from other areas. One such 
plate is that forming the Chu-Ili Range, and termed here the Chu-Ili 
Plate (Fig. 1 ). Within the Chu-Ili Plate the successions have been 
known for some time (e.g. Nikitin 1972, 1973). However, although 
a number of papers have been published on aspects of some of the 
contained Ordovician faunas, much remains to be done. A central 
formation within the unit is the Anderken Formation of early Caradoc 
age. This immediately underlies the Dulankara Formation, whose 
brachiopods from its lowest Otar Member we have recently revised 
( Popov et al. 2000). Although some pioneering descriptions of some 
of the Anderken brachiopods were published by Rukavishnikova 
( 1956) and some individual species have been published in a number 
of publications, e.g. Popov (1980, 1985) and Nikitin & Popov 
( 1983), the whole brachiopod fauna from the formation has never 
been published, and this is the chief purpose of the present paper. In 
addition, six brachiopod-dominated associations can be identified 
from the Anderken Formation. LEP and LRMC are responsible for 
the whole paper and IFN for input into the systematic palaeontology 
and biofacies sections. 



OUTLINE OF GEOLOGY AND FOSSIL 
LOCALITIES 

The Chu-Ili Plate (Fig. 1), as recognised here, is a small part of Asia 
today, and is traceable from the Zailiyskiy Alatau Range in the 
southeast to the northern Betpak-Dala Desert in the northwest, 
where it disappears under late Palaeozoic and Mezo-Cenozoic de- 
posits. To the southwest it is bordered by the large Dzhalair-Najman 
Fault and northward-dipping homoclinal sequences of Upper 
Cambrian and Lower to Middle Ordovician age which are mainly 
siliciclastic slope rise deposits (e.g. the Dzhambul Formation), indi- 
cating passive margin development, and several thrust sheets 
consisting of dismembered ophiolites of the early Palaeozoic 
Ashchisu Formation (Toporova et al. 1971). The Dzhalair-Najman 
Fault mainly follows an early Palaeozoic suture which separates the 
Lower Palaeozoic Chu-Ili Plate from the Middle to Upper Ordovician 
volcanic island-arc association traceable along the northeastern 
margin of the Betpak-Dala-North Tien Shan tectonofacies belt of 
Nikitin (Nikitin et al. 1991), which is the same as the Djezkazaan- 
Kirgiz (4.1) tectonofacies unit of Sengor & Natalin (1996). To the 



UPPER ORDOVICIAN BRACHIOPODS FROM KAZAKHSTAN 



15 






Chu-lli Plate 



Zhalair-Najman Synclinorium 



Central pari 



Southern part 



Burultas tectonofacies belt 



Assemblage of 
fore-arc basin and 
onlap assemblage 
(Sarytuma Zone) 



Subduction-accretion complex 



Burubaital Package 

(dismembered 

ophlollte) 



Maikul Packege 

(slope-rise 

deposits) 



Zhalgyz Packege 

(dtsmembered 

ophlollte) 



Darbaza Packege 
(carbonate platform) 



Zhalalr 
Formation 



Zhalair Formation 

(siiistone and mudstone, interlayers of limestone and tuff) 



Chokpar 



Chokpar 
Formation 



Ulkuntas 
Limestone 



> 



Chokpar Formation 

(siltstone and mudstone, bla 



Kyzylsai 
Formation 



Kyzylsai Formation 

(sandstone, polymlct 
and siltstone) 



Dulankara Formation 

(conglomerate polymict, sandstone, siltstone, lenses of limestone) 



Anderken Formation 

(conglomerate pofymlct. sandstone, siltstone and limestone) 



Kopaly 



Baigara Fm. 

(limestone, 
sandstone and 
siltstone) 

Ka ratal 
Formation 

(sandstone, 
siltstone. 
mudstone, and 
tuff) 



Kogashik 



Aktau 



Ungur 



Dzhambul 
Formation 

(sandstone, 
quartzose, 
siltstone and 
mudstone, 
graded) 



Beke Formation 

(sandstone, siltstone 
and mudstone, graded) 



Uzunbulak Formation 

(sandstone, siltstone and 
mudstone, interlayers of 
limestone, mass flow 
deposits) 

"Akzhal" Formation 

(limestone, bedded, 
conglomerate and 
sandstone at the base) 



PrecambHan basement 



Unnamed formation of 
sandstone and graded 
siltstone. with 
olistostrome horizon 
at the base 

Oisaksaul Formation 

(conglomerate, sandstone, 
siltstone and limestone) 



Alakul 
Bolgozha / Limestone 
Formation 

(siliceous shales, black, 
siliceous; extrusives 
and tuff, rhyolite-dacite. 
mass flow deposits) 



Burubaital 
Formation 

(black and red 

radiolarian 

chert) 




Maikul 
Formation 

(sandstone, 
quartzose. 
siltstone and 
mudstone, 
graded) 



Zhalgyz 
Formation 

(sandstone, 
olcano-clastic and 
siltstone, graded 
black shale, 
basaltic extrusives 
and tuff) 



Darbaza Formation 

(quartzes 
limestone 
dolomite) 

niDiroiii 



Angular unconformity 



Discontinuity 



q Stratigraphic contact 
unknown 



Fig. 2 Chart showing the correlation between the Lower Palaeozoic lithostratigraphic units in the Chu-lli Plate and the Burultas tectonofacies belt. 



north-east of the Chu-lli Plate lies the Burultas tectonofacies belt 
(Fig. 2), which represents an accretionary wedge suggesting active 
margin development, island arc volcanism and subduction of the 
oceanic crust under the Chu-lli Plate from the early Arenig to the 
Llandeilo Pygodus anserinus Biozone (Koren et al. 1993). By the 
Caradoc, subduction and volcanism had ceased as a result of the 
docking of a small terrane or island arc, the Mynaral-South 
Dzhungaria tectonofacies belt of Nikitin et al. ( 1 99 1 ). 

The Ordovician deposits in the central part of the Chu-lli Plate, the 
Dzhalair-Najman Synclinorium, form a nearly continuous sequence 
of siliciclastic and carbonate rocks from Arenig to Ashgill in age 
(Fig. 2), which are relatively unmetamorphosed and chiefly dip 
gently to the northeast. They are covered conformably by Silurian 
deposits (Nikitin et al. 1980) or unconformably by the Devonian. 
The Ordovician stratigraphy and major lithostratigraphic units were 
described by Keller (1956) and Nikitin (1972). 

The Lower to Middle Caradoc deposits, which are the main source 
of the brachiopods described here, are termed the Anderken Forma- 
tion, which is a transgressive sequence of mainly siliciclastic deposits 
that contain variably developed lens-like carbonate units in the upper 
part representing mud mounds or algal build-ups (Nikitin et al. 1974; 
1996). They are best developed in the following eight general 
localities (Figs 1,3): 

Localites 1-2. Area between the Ashchisu and 
Sarybulak rivers. 

In the south-eastern part of the Chu-lli Range the best sections of the 
Anderken Formation are located in a block with faulted margins 
between the rivers Ashchisu and Sarybulak (Fig. 1, localities 1-2; 



Figs 3, 4). Here the Anderken Formation overlies, with a slight 
angular unconformity, graded sandstones, siltstones and mudstones 
of the Beke Formation, which is Llandeilo to early Caradoc in age, 
dated by numerous graptolites of the Hustedograptus teretiusculus 
and Nemagraptus gracilis Biozones (Tsai 1976). The Anderken 
comprises six lithostratigraphic units traceable up to 40 km along 
strike, overlain unconformably by Devonian deposits. These units 
are (in ascending order): 

Unit 1. Polymict, pebbly conglomerate, with sandy matrix and with 
some beds of sandstone and gritstone. Thickness from 45 m to 120 
m, with maximum values in the Anderkenyn-Akchoku section. 

Unit 2. Coarse- to medium-grained sandstone with subhorizontal 
stratification alternating with abundant cross-bedded sets. Lenses of 
polymict, pebbly conglomerate represent shallow channels formed 
by tidal currents. Thickness varying from 52 m in the Kujandysai 
section in the west to 180 m in the Anderkenyn-Akchoku section 
(Figs 3, 5). The upper part contains the lingulide Ectenoglossa 
sorbulakensis, the trilobite "Isotelus" romanovskyi Weber and gas- 
tropods (Samples 8130-1, 7612). The middle part of the unit in the 
Anderkenyn-Akchoku section contains a carbonate mud-mound up 
to 16 m thick with a core built of light grey micritic limestone. On the 
flanks there is bedded biomicrite with the brachiopods Skenidioides 
sp., Christiania sp. and Kellerella misiusi, the trilobites 
Mesotaphraspis spinosus Lisagor, Selenoharpes sp., Acrolichas sp. 
Eokosovopeltis romanovskyi and Sphaerexochus aff. hisingeri War- 
burg. Illaenus sp. was noted from about 1 .0-1 .5 m below the top of 
the mud-mound in the eastern part of its exposure (Sample 8226). 
Cystoid and crinoid columnals are abundant. 



16 



L.E. POPOV, L.R.M. COCKS AND I.F. NIKITIN 



Devonian 



1 . Anderkenyn-Akchoku 
section (west) 
l-»- 2. Kujandysai section 



4. Buidukbai-Akchoku 
Mountain 




8. Kotnak mountains 
SWof Karpkuduk 



Devonian 

7. South side of 
Karatal river south 
of Sorbulak spring 300 



6. Burultas 
valley 



A.-D.A. ,oo 
F-1041a 
«■ 818a 
T.A. 

50 



2 



F-1024b 
T.A. , 



F-1024 
Ect. A. 



T.A. 

.-F-1018 

e-F-1018a 

T.A. 



Conglomerate and sandstone, 
intercalating 



Intercalating sandstone 
and siltstone with coquina 
storm beds 

Limestone and siltstone 
intercalating 

Sandstone, bedded. 

Sandstone, cross-bedded 

Sandstone 



7*TT71 Conglomerate, pebbly, 
polymict 



Beke Formation 






Sandstone, siltstone 
and mudstone, graded 



MIDDLE ORDOVICIAN 
Uzunbulak Formation 



Siltstone and mudstone 



Fig. 3 Columnar sections through the Anderken Formation showing informal units, stratigraphic positions of samples and brachiopod associations: Ect. A.- 
Ectenoglossa Association, T. A., Tesikella Association, M.S. A.. Mabella-Sowerbyella Association. A.-D. A., Acculina-Dulankaretla Association, P.-K. A., 
Parastrophina-Kellerella Association, Z.-F. A., Zhilgyzambonites-Foliomena Association, G.-B. A, Gastropod-Bivalved Molluscs Association. The 
numbers of the sections are the same as those on Fig. 1 . 



Unit 3. Coarse- to medium- grained sandstone with mostly 
subhorizontal stratification, about 40-62 m thick in the Anderkenyn- 
Akchoku section and up to 97 m thick in the Kujandysai section, with 
some storm beds of coquinas up to 20 cm thick with concentrations 
of gastropods and the disarticulated bivalved molluscs Edmondia 
fecunda Khalfin, Ctenodonta sp. and Orthonota'l sp. (Sample 8 1 30a). 
Gastropods, bivalved molluscs and the trilobite Eokosovopeltis 
romanovskyi become increasingly abundant in the flank deposits in 
the upper 20 m of the unit (Samples 8130, 8134). In the Kujandysai 
section concentrations of bivalved molluscs occur in the middle part 
in association with the rare brachiopod Tesikella necopina and the 
pelmatozoan columnals Clivosocystis clivosus Stukalina and 
Ordinacrinus punctatus Stukalina (Sample 761 1). 

Unit 4. Medium- to fine-grained sandstone replaced gradually up- 
wards by siltstone with numerous trace fossils and symmetrical 
ripple marks. Thickness varies from 22 to 80 m in the Anderkenyn- 
Akchoku section and is about 28 m in the Kujandysai section. The 
lower part contains local concentrations of the coalified plant 
Akdalaphyton caradoci Senkevich, and gastropod and bivalved mol- 



luscs in association with the brachiopod Tesikella necopina (Sam- 
ples 8 127-2b, 8 1 29, 8 1 33, 8 1 38). The upper part contains an abundant 
brachiopod fauna of the Sowerbyella-Mabella Association (Sam- 
ples 100b, 7613. 8128a, 8128b, 8135, 8137) and the trilobites 
Dulanaspis laevis anderkensis Chugaeva, Lonchodomas tecturmasi 
Weber, Pliomerina sp., Remopleurides sp., Styginella macrophtalma 
Pribyl & Vanek, Bronteopsis extraordinaris Chugaeva, the cystoid 
and crinoid columnals Clivosocystis clivosus, Digiticrinus levis 
Stukalina, Ordinaricrinus punctatus Stukalina and Communicrinus 
communis Stukalina and the starfish Stenaster obtusus (Forbes). 

Unit 5. Limestones varying in thickness from 8 to 98 m forming a 
chain of carbonate build-ups between the Uzunbulak and the Ashchisu 
rivers (Fig. 3). The cores of these build-ups rest on a bed of nodular 
limestone from 2-10 m thick with abundant dasyclad algae 
Cyclocrinites nikitini Gnilovskaya and Mastopora reticulata 
Gnilovskaya (Nikitin etal. 1974). A bed of nodular, algal limestone 
with dasyclad algae is usually present in the interspaces between the 
carbonate build-ups and contains brachiopods of the Acculina- 
Dulankarella Association (Samples 100, 8251. 85258), the rare 



UPPER ORDOVICIAN BRACHIOPODS FROM KAZAKHSTAN 



17 



Conglomerate and 
red sandstone 



MIDDLE - UPPER ORDOVICIAN 
Anderken Formation 



Siltstone and 
mudstone (Unit 6) 

Limestone 
(Unit 5) 

Sandstone and 
siltstone (Unit 4) 



I | Sandstone 



(Unit 2-3) 

Conglomerate, 
polymict (Unit 1) 



Beke Formation 

Sandstoi 
mudstone, graded 



| : x : : : : : :| Sandstone, siltstone and 




Fig. 4 Geological map showing distribution of the Middle and Upper Ordovician rocks and the positions of measured sections and fossil localities that 
yielded brachiopods in the area between the Uzunbulak and Ashchisu Rivers, south-eastern Chu-Ili Range (after Nikitin 1972, modified). 



tabulate corals Lichenarial sp. and Amsassia sp., stromatoporoids, 
and various trilobites and echinoderms. Locally between the 
Kujandysai and Sarybulak rivers, and on both sides of the Ashchisu 
River in the eastern part of outcrop area, carbonate build-ups disap- 
pear and the unit comprises bedded limestone varying from biomicrite 
to biosparite intercalating with siltstone and mudstone, with 
brachiopods of the Parastrophina-Kellerella Association (Tables 4— 
5, Samples 628, 8223, 8223a, 8223b). Brachiopods of this association 
also occur in pockets of bioclastic limestone in the mud-mound core 
and the overlying bedded limestone together with large spherical or 
ellipsoidal ooids of radiaxial calcite up to 1 cm across (Samples 
2538,8217,8219,8256). 

Unit 6. Siltstone and mudstone with up to 6 interlayers of bentonite 
up to 0.3 m thick in the lower part, total about 50-60 m thick, 
containing brachiopods of the Zhilgyzambonites-Foliomena Asso- 
ciation (Samples 8231, 8251, 8255). Abundant trilobites are 
Gramilatagnostus granulatus Kolobova, Sphaeragnostus sp., 
Microparia speciosa Hawle & Corda, Hammatocnemis sp.. 
Birmanites almatiensis (Chugaeva), Cyclopyge sp., Cybele weberi 
Chugaeva and Ovalocephalus sp., and graptolites include 
Dicranograptus nicholsoni, Diplograptus anderkenensis, 
Glyptograptus trubinensis and Pseudoclimacograptus scharenbergi, 
suggesting the Lower to Middle Caradoc Diplograptus midtidens 
Biozone (Keller 1956). 

Locality 3. East side of Kopalysai River 

On the east side of the Kopalysai River (Fig. 1) the Anderken 
Formation is about 160 m thick and rests unconformably on the 
siliciclastic Llandeilo Beke Formation (Fig. 2). Detailed description 
of this section was provided by Keller (1956: 26), who recognised 
three units (Fig. 3): (1) bed of intercalating polymict conglomerate 



and coarse- to medium-grained sandstone up to 70 m thick with 
bivalves, rare Ectenoglossa sorbidakensis (Sample 8223-1) and 
numerous plant remains of Akdalaphyton caradoci; (2) intercalating 
fine-grained sandstone and siltstone about 15-20 m thick with 
brachiopods of the Tesikella Association (Sample 1 27), the trilobites 
Duhmaspis levis anderkensis and Lonchodomas tecturmasi; and (3) 
mudstones with some siltstones and fine-grained sandstones about 
70-80 m thick with abundant brachiopods of the Mabella- 
Sowerbyella Association (Sample 8228). The deposits overlying the 
Anderken Formation are polymict pebbly conglomerates and 
sandstones of the Dulankara Formation. 

Locality 4. Buldukbai-Akchoku Mountain 

On the west side of the River Kopalysai, the Anderken Formation 
includes a large carbonate mud-mound which forms the top of 
Buldukbai-Akchoku Mountain. The lower part of the formation is 
exposed on the south-western slope of the mountain, north of an east- 
west fault (Figs 1, 3, 5-7). It includes, in ascending order: 

Unit 1 . Medium- to fine-grained sandstone up to 120 m thick with 
Ectenoglossa sorbidakensis. 

Unit 2. Dark green, bedded siltstones, about 14 m thick with a few 
layers of fine grained sandstone 3-10 cm thick, containing Mabella 
conferta and Shlyginia fragilis of the Sowerbyella-Mabella Asso- 
ciation. 

Unit 3. Siltstones with nodules of algal limestone gradually chang- 
ing into beds of nodular limestone with dasyclad algae towards the 
top, 38 m thick in total. 

Unit 4. Nodular algal limestone intercalating with siltstone about 
0.5-1.5 m thick, up to 22 m thick in total, with brachiopods of the 
Acculina-Dulankarella Association (Sample 8231-40). 



18 



L.E. POPOV, L.R.M. COCKS AND I.F. NIKITIN 



Anderkenyn-Akchoku Section (eastern side) 



Parastrophina 

Kellerella 

Association 



Ectenoglossa 
Association 
8134 



Tesikella 
Association 

Bivalved 8134a 

molluscs and 
gastropods 

8134b 




Ma bell a ■ 
Sowerbyella 
Association 
8235 8137 621 

Unit 5 



Uunit 5 



\ 50 m 
Unit 2 \/ Unit 3 

' 25 m N 



55 m 



Kujandysai Section 



Tesikella 
Association 

7611 



Parastrophina - Kellerella Association 
2538 



Mabella ■ Sowerbyella 
Association 



Ectenoglossa 
Association 






Unit 1 



52 m 
Unit 2 






8219 




Unit 6 



Buldukbai-Akchoku Section 




Acculina - Dulankarella 
Association 



Ectenoglossa 
Association 



Mabella - Sowerbyella 8231 ■ 
. Association 

110 



120 m 




8227-80 

8227-40 
8227-10 



Units 



36 m ""* Unit 4 
\/"\/ Unit 3 

Unit 2 I 

Fault 



Mabella - Sowerbyella Association 



Unit 1 
20 40 60 80 100 




Unit 6 



DEVONIAN 

Conglomerate 



UPPER ORDOVICIAN 
Dulankara Formation 



>*•*•* Conglomerate, pebbly, polymict 



Anderken Formation 

I I Siltstone and mudstone 



oa 



Limestone, massive 
micritic 



pep|pe|| Limestone, algal, 
:g^$<a nodular 



Siltstone 



-- vr^jH Intercalating sandstone 
r -r-:-r-:-T:'-r and siltstone 



Sandstone, 
cross-bedded 



— -^ Sandstone, bedded. 
Sandstone 



252 J Conglomerate, pebbly, 

J oVoVoj polymict 



Fig. 5 Schematic stratigraphic sections of the Anderken Formation in the Chu-Ili Range, showing the position of samples and distribution of brachiopod 
associations. 



UPPER ORDOVICIAN BRACHIOPODS FROM KAZAKHSTAN 



19 



626 8223a, 8223b 



843 



100b 8221 A 

Devonian 




Fig. 6 A, general view of the Anderken Formation at the Anderkenyn-Akchoku section, showing informal lithostratigraphic units discussed in the text and 
the position of brachiopod localities. B, view of large complex carbonate buildup at Akchoku Mountain in the upper part of the Kujandysai section. 
Photographs by Igor Nikitin. 



Unit 5. Massive micritic limestone forming the core of the carbon- 
ate mud-mound at the top of the mountain, about 30 m thick. 

The upper part of the Anderken Formation outcrops along the 
north-eastern slope of Buldukbai-Akchoku Mountain. It includes: 

Unit 6. Laminated, dark green siltstone up to 70 m thick with storm 
beds of calcareous sandstone rich in brachiopod coquinas about 10- 
15 cms thick and crinoid columnals. The top is a characteristic bed of 
laminated brownish-violet siltstone about 0.5 m thick overlain by 
polymict conglomerate of the Dulankara Formation (Fig. 7B). The 



unit contains numerous coalified plant remains of Akdalaphyton 
caradoci concentrated on several bedding surfaces, brachiopods of 
the Sowerbyella-Mabella Association (Samples 8229, 8230) and the 
echinoderms Clivosocystis sp., Digitocrinus levis and Ristnacrinus 
bifidus Stukalina. 

Locality 5. Tesik River 

This locality is on the southern side of the River Tesik about 1.5 km 
upstream of the bridge crossing the river on the highway from 



20 



L.E. POPOV, L.R.M. COCKS AND I.F. NIKITIN 



Unit 5 



8231-40 














."••in 










Fig. 7 The Anderken Formation on the south-western slope of Buldukbai-Akchoku Mountain. A, Units 1 to 5, and Sample localities 8227-80, 1 10 and 
8231-8240; B, Unit 6 and the contact with the overlying Dulankara Formation and Sample localities 8229 and 8230. Photographs by Lars Holmer. 



Almaty to Balkhash (Fig. 1, locality 5). It is an isolated natural 
exposure of about 20 m of pink to light red rocks of massive 
micritic limestone forming the core of a mud-mound with lens- 
like beds of biosparite at the base of the exposure. Most of the 
bioclasts are fragmented large cystoid columnals. It contains an 



abundant Parastrophina-Kellerella Association (Sample 948). 

Locality 6. Burultas Valley 

The Burultas Valley (Fig. 1, locality 6) is about 42-45 km west of 
Chiganak on the western Balkhash coast (northeastern part of 



UPPER ORDOVICIAN BRACHIOPODS FROM KAZAKHSTAN 



21 



Quadrangle 73°22'30" to 73°30' E; 45° to 45°05' N). A summary of 
the Ordovician geology and lithostratigraphy of this locality is in 
Nikitin et al. (1980, text-figs 18, 20). The Anderken Formation 
consists mainly of siliciclastic rocks with a thick unit of polymict 
conglomerates at the base and a number of carbonate mud-mounds 
in the upper part (Fig. 3). The carbonate unit in the top of the 
sequence is a bed of nodular algal limestone about 6- 1 m thick with 
numerous Girvanella sp., Cyclocrinites nikitini and Mastopora 
reticulata and brachiopods of the Acculina—Dulankarella Associa- 
tion (Locality 1041a of Nikitin = Sample 390/76 of Kovalevskii). 
which underlies a lens of massive, micritic limestone up to 20 m 
thick which forming the mud-mound core. The unit thins about 200 
m westward from Locality 1041 a, where it is represented by bedded 
and nodular limestone with the brachiopods Pionodema opima, 
Dulankarella larga and Mabella conferta (Sample 818). The upper- 
most 10 m of the underlying unit, of fine-grained sandstone 
intercalating with siltstone, contains a different brachiopod assem- 
blage with Tesikella necopina (Sample 818a), in association with 
abundant cystoid columnals. 

Locality 7. Sorbulak spring on the east side of the 
River Karatal 

In the south Betpak-Dala Desert, about 20 km west of Baigara 
Mountain, the Anderken Formation is well exposed on both sides of 



the River Karatal (Fig. 1, locality 7). Here it rests on the graded 
sandstones and siltstones of the Llandeilo to Lower Caradoc upper 
Baigara Formation (Fig. 2), or is in contact with intrusives. About 2 
km south-east of the Karatal river, south of Sorbulak spring, it 
comprises (1) polymict conglomerates more than 50 m thick, (2) 
medium- to fine-grained sandstones 169-170 m thick with 
Ectenoglossa sorbulakensis about 10-15 m above the base of the 
unit (Fig. 3, Sample 1024); and (3) intercalating fine-grained slightly 
calcareous sandstones and siltstones about 60 m thick with the 
Tesikella Association in the upper 20 m of the unit. The upper part of 
the section is an unfossiliferous unit of intercalating fine-grained 
sandstones, lilac and red siltstones and mudstones several hundred 
metres thick, which is overlain by the basal conglomerate of the 
Dulankara Formation. 

Locality 8. Kotnak Mountains 

This incomplete section of the Anderken Formation is situated west 
of the Kotnak Mountains, about 7 km SW of Karpkuduk Well. There, 
about 1 .5 km north-east of the salt marsh (Figs 1 , 3, 8), the formation 
consists of: ( 1 ) siltstone about 70 m thick with some storm beds of 
calcareous sandstone about 10-20 cm thick with a coquina of the 
bivalve Ctenodonta sp. (Samples 1017, 1019); (2) sandstone inter- 
calating with siltstone in the upper part, total 40 m thick, with 
brachiopods of the Tesikella Association in the lower 10 m of the unit 



QUATERNARY 



Salt marsh 



DEVONIAN 



ES 



ORDOVICIAN 
Anderken Formation 



Conglomerate 



Sandstone 



vv^j Sandstone and 
t>;r i£3 siltstone 



*':X 





Sandstone with coquina 
storm beds 





i i_ 



2 km 



Fig. 8 Geological map showing the distribution of the Anderken Formation and the position of fossil localities in the area about 7 km south-west of 
Karpkuduk well, Kotnak Mountains. 



22 



L.E. POPOV, L.R.M. COCKS AND I.F. NIKITIN 



Table 1 Composition of Ectenoglossa Association from the Anderken Formation showing number of complete shells, ventral and dorsal valves 
respectively. 



Sample number 
Number of specimens 
Ectenoglossa sorbulakensis 



7612 


8130a 


8130- 


27 


1 


2 


24:3:2 


0:1:0 


0:2:0 



8223-1 



0:1: 



8227-10 


8227^10 


8227-80 


F-1024 


6 


4 


3 


12 


6:0:0 


0:3:4 


0:3:3 


0:8:12 



(Sample 1018a) below a bed of skeletal calcareous sandstone about 
5 m thick with an allochthonous brachiopod fauna with a mixture of 
taxa of the Tesikella and Mabella-Sowerbyella Associations (Sam- 
ple 1018); (3) intercalating beds of sandstone and pebbly polymict 
conglomerate about 80-90 m thick; and (4) siltstone with a few beds 
of fine-grained sandstone, total 130 m thick and overlain 
unconformably by Devonian conglomerate. 



FAUNAL ASSOCIATIONS 

The Anderken Formation is a transgressive sequence from near shore 
to outer shelf deposits with predominantly siliciclastic deposition. 
The lower part of the formation, below the main horizon with 
carbonate build-ups in theAnderkenyn-Akchoku, Kujandysai, 
Buldukbai-Akchoku and Burultas sections, was formed in tide- 
dominated environments of mostly tidal flat deposits with 
characteristic sets of pebbly conglomerates, cross-bedded and lami- 
nated sands, coquina storm beds and traces of tidal currents. Carbonate 
build-ups in the upper part of the formation preserve numerous 
traces of photosynthetic activity and contain a diverse flora of green 
and red algae (Nikitin et al. 1974); suggesting formation in shallow 
depths within the euphotic zone. The outer shelf deposits are recorded 
only in the south-eastern Chu-Ili Range and consist of silt and mud 
containing graptolites. The benthic fauna is dominated by trilobites 
but includes one of the earliest records of the Foliomena brachiopod 
fauna. Apollonov (1975) has described the trilobite associations of 
the middle and late Ordovician of the Chu-Ili Plate. 

A matrix based on the distribution of about 1800 brachiopod 
specimens from 33 samples within the Anderken Formation was 
subjected to Principal Component Analysis (Etter 1999). Plots of 



eigenvectors corresponding to three maximum directions of varia- 
tion (F1-F3) are illustrated on two two-dimensional diagrams (Fig. 
9). The Diversity Index is calculated as the number of species minus 
1 divided by the natural logarithm of the number of brachiopod 
individuals in the sample (for details see Williams et al. 1981 ).The 
analysis of taxonomic composition and relative abundance of 
brachiopod taxa from numerous localities and samples in the 
Anderken Formation allows recognition of six brachiopod associa- 
tions characterised below. They are interpreted within the Benthic 
Assemblage (BA) scheme of Boucot (1975). 

1. The Ectenoglossa Association. This is a monospecific lingulide 
association of BA-1 with Ectenoglossa sorbulakensis in the 
Anderkenyn-Akchoku, Kujandysai and Buldukbai-Akchoku sec- 
tions, the east side of the Kopalysai River and on the southern side of 
the Karatal River south of Sorbulak spring (Table 1 ).The assemblage 
shows patchy distribution in lithologies of coarse- to medium- 
grained sands with subhorizontal and cross-bedded stratification. In 
most of the localities shells are disarticulated on the bedding surfaces 
and only in Sample 7612 do conjoined valves predominate (89% of 
individuals). A cluster of six articulated shells preserved in life 
position inclined from 62°-80° to the bedding surface was recovered 
from Sample 8227-40 in the Buldukbai-Akchoku section, which 
confirms the infaunal mode of life of this lingulide. The gastropods 
Lophospira sp. and Latitenia kasachstanica Vostokova and the 
bivalved molluscs Endomionia fecinda, Ctenodonta sp. and 
Cyrtodontal subcentralis (Khalfin 1958), which are widespread in 
similar lithologies and form coquina storm beds, do not co-occur 
together with the lingulides; for example, in Sample 8 1 30 a bedding 
surface with Ectenoglossa sorbulakensis and a storm bed with 
molluscs and the trilobite Eokosovopeltis romanovskyi are separated 
by an interval only about 2.5 m thick. It is likely that Ectenoglossa 



Table 2 Composition of Tesikella Association from the Anderken Formation showing number of complete shells, ventral and dorsal valves respectively. 



Sample numbers 


127 


818a 


F-1018a 


7611 


8128 


F-1018 


F- 1024b 


Number of specimens 


18 


22 


32 


7 


18 


172 


20 


Diversity index 


0.33 


0.65 


0.58 


0.51 


1.38 


3.11 


2.16 


Trematis sp. 


0:0:1 














Tesikella necopina 


3:14:12 


0:13:12 


0:16:7 


1:5:5 


0:4:2 


1:4:3 


0:2:2 


Longvillia lanx 






0:4:5 






2:7:3 




Glyptomena onerosa 












0:1:5 




Christiania egregia 












0:7:18 


0:2:7 


Limbimurina sp. 












0:1:1 




Isophragma imperator 












1:36:34 




Acculina kulanketpesica 










0:0:1 


0:1:2 




Mabella conferta 




1:0:0 








0:1:2 




Shlyginiafragilis 












0:6:4 




Anaptambonites orientalis 












1:4:3 




Sowerbyella rukavishnikovae 






0:10:11 






0:8:17 


0:9:10 


Bicuspina rukavishnikovae 












3:1:9 




Plectorthis bundtasica 










0:0:1 


1:6:7 




Dolerorthis expressa 












1:12:9 




Phragmorthis conciliata 












0:4:4 


0:1:0 


Eodalmanella extera 




4:4:0 




0:0:1 




1:28:22 




Pionodema opima 










2:8:5 






Rhynchotrema sp. nov. 










1:0:0 






Didxmelasma cf. transversa 












0:0:1 





8127-2b 



8138 
2 



0:1:0 



0:2:0 



UPPER ORDOVICIAN BRACHIOPODS FROM KAZAKHSTAN 



23 



F2 



• 2538 



Parastrophina - Kellerella 
Association 

100 + *8217 
/ \ -k 8256 
626 / •• 626 

T, 8214 \&»8 * 

Acculina - Dulankarella 's* \* . 8226 
Association \„ £«223b 

622 ^-* \ B219 

8223* \* 
6220 . 
8215-». w 390 
85258 X »| U ~V 
- 1.0 -0.8 -0.6 -0.4 e ; 3 ,T^ 



Zhilgyzambonites - X31 * 
Foliomena Association 8255 



F3 

0.6 



Zhilgyzambonites - 8J55 
Foliomena Association ,„.'< 



Mabella - Sowerbyella 
Association 

8128b 8257 8 228 



Acculina - Dulankarella 
Association 

626 . .,. '" 

T" -0.4 



8135 

0.2* 



100b 
0.6 



F1 

1.0 



8128 
2 
F-1024b" 



V7611 
V 



F-1018 
V F-1018a 

Tesikella Association 



J5258 

)1^oT+ 



— 



8223 



• 8215 
8220 

• 



\ 

100 4, 

82233^628 
8223b ■& 
8214* * 

8219 

8256 i, * B 226 1 

8217* * 
948 

Parastrophina - Kellerella 
Association 



8135 . 

- 0.4 

8251 

- 0.2 
390 



Mabella - Sowerbyella 
Association 

F1 

0.4 m" 0.6 0.8 1 .0 



F-1024b 

--0.2 



F-1018 Y 7613 • B137IJ 

6230 ■ B 843" - 
8128b ,„,.8128a 



8157 
8228 



▼ 

T 

8128 



Tesikella Association 



Biuay 
-0.6 F-1018a 



Fig. 9 Two-dimensional principal component analysis plots on first (Fl ). second (F2) and third (F3) eigenvectors of selected brachiopod samples from the 
Anderken Formation shown on Tables 1-6. 



sorbulakensis formed a separate monotaxic community which in- 
habited a mobile sandy bottom in peritidal environments, perhaps 
tidal flats. 

2. The Tesikella Association (average Diversity Index 1.18; observed 
range 0.33-2. 16, N=7). This is a low-diversity strophomenide-domi- 
nated brachiopod association. It is widespread in shallow marine 
environments of BA-2, which are mainly fine-grained sands with 
subhorizontal stratification, occasional storm beds with mollusc 
coquinas and locally abundant plant remains of Akdalaphyton 
caradoci. This association is defined by the index species Tesikella 
necopina, an endemic species and genus restricted only to the Lower 
Anderken Formation. This species pursued an opportunistic life 
strategy and expanded into environments inhabited mostly by gas- 
tropod and bivalved mollusc communities, in which 
rhynchonelliformean brachiopods, if they occur, are an insignificant 
component of the assemblage and show patchy distribution. Traces 
of tidal currents, ripple marks and occasional storm beds suggest 
rather turbulent environments occasionally affected by seasonal 
storms. Shells in all the samples are mostly disarticulated (Table 2) 
and some contamination by allochthonous shells from adjacent 
associations cannot be excluded. However, results of the Principal 
Component Analysis show that all the samples referred to the 
Tesikella Association form a distinct cluster and are characterised by 
low positive values of Fl and negative values of the two other 
maximum directions of variation (F2 and F3). Sample 1018, which 
may be contaminated by allochthonous shells, shows low negative 
values of F3 similar to the samples of the Mabel la-Sowerbyella 
Association (Fig. 9). 

In its pioneer stage, the Tesikella Association is characterised by 
the appearance of Tesikella necopina in mollusc-dominated environ- 
ments, where it is the only brachiopod ( Samples 127,7611,81 27-2b, 
8138), or where it comprises more than 50% of the brachiopod fauna 
together with Longvillia lanx, Sowerbyella nikavishnikovae (up to 
34%) and Eodalmanella extern (up to 18%). At its mature stage the 
association includes four to eight species usually common in the 
Sowerbyella-Mabella Association, e.g. Christiania egregia (up to 
35%), Sowerbyella nikavishnikovae (up to 50%) and Pionodema 
opima (56% in one sample). All the other taxa constitute less than 5% 
of individuals in any particular sample. Among other groups bivalved 
molluscs, the trilobites "Isotelus" romanovskyi, Lonchodomas 



tecturmasi and Dulanaspis levis anderkensis occur. The echinoderm 
fauna is dominated by the cystoidean Clivosocystis minusculus 
Stukalina which is known only from columnals. The abundance of 
coquinas and plant remains suggests biogenic fixation of a sandy 
substrate. 

Sample 1018 from the Kotnak Mountains is placed within the field 
of the Tesikella Association (Fig. 9), but differs in high taxonomic 
diversity and the occurrence of taxa characteristic of the Sowerbyella- 
Mabella and Acculina-Dulankarella associations, e.g. 
Anoptambonites orientalis, Glyptomena onerosa, Limbimurina sp. 
and Acculina kulanketpesica. This sample came from a bed of 
bioclastic sandy limestone, which is an atypical lithology for the 
Tesikella Association, more likely to have been deposited within a 
bar system, and contains an allochthonous brachiopod coquina 
representing a mixture of several life associations. The abundance of 
coarse elastics and storm beds formed mostly by the bivalve 
Ctenodonta sp. (Samples 1017, 1019) in the Anderken Formation of 
this section suggest turbulent depositional environments within a 
shore-face zone comparable with the lower part of the Otar beds in 
the Dulankara Formation of the Dulankara section in the south- 
eastern Chu-Ili Range (Popov etal. 2000). 

3. Mabella-Sowerbyella Association (average Diversity Index 1 .30; 
observed range 0.83-1 .86, N=9) is another low diversity association 
of BA-2 dominated by strophomenides. It is recognised in the 
Anderkenyn-Akchoku, Kopalysai and Buldukbai-Akchoku sections 
by the predominance of Mabella conferta and Sowerbyella 
nikavishnikovae which together comprise 40-80% of individuals in 
the assemblage. Glyptomena onerosa, Shlyginia fragilis and 
Anoptambonites orientalis mainly occur in this association (Table 
3). As in the mature Tesikella Association, orthides are represented 
only by Eodalmanella extera and Pionodema opima, which do not 
usually co-occur. This association is confined to a fine clastic substrate 
of silts and fine-grained sands, usually with traces of bioturbation 
and locally abundant concentrations of the plant Akdalaphyton 
caradoci. In samples from the Anderkenyn-Akchoku section 
brachiopods are preserved disarticulated, but in the Kopalysai and 
Buldukbai-Akchoku sections the number of articulated specimens 
increases up to 50 percent, suggesting rapid burial and lack of 
significant post-mortem displacement of the shells. The most com- 
mon trilobites in the associated faunal assemblage are Lonchodomas 



24 



L.E. POPOV, L.R.M. COCKS AND I.F. NIKITIN 

valves 



Table 3 Composition of Mabella-Sowerbyella Association from the Anderken Formation showing number of complete shells, ventral and dorsal 
respectively. 



Sample number 


100b 


7613 


8128a 


8128b 


8137 


8257 


8228 


8229 


110 


8230 


843 


Number of individuals 


248 


50 


9 


27 


10 


21 


41 


5 


11 


36 


75 


Diversity index 


1.09 


1.79 


1.82 


1.21 


1.73 


1.31 


1.62 


1.86 


0.83 


1.13 


1.39 



Trematis sp. 
Paracraniops sp. 
Glyptomena onerosa 
Christiania egregia 
Foliomena prisca 
Dulankarella larga 
Mabella conferta 
Shlyginiafragilis 
Anoptambonites oriental is 
Olgambonites insolita 
Sowerbyella rukavishnikovae 
Triplesia sp. 

Phragmorthis conciliate! 
Plectorthis burultasica 
Eodalmanella extera 
Pionodema opima 
Rhxnchotrema akchokense 



0:0:1 



0:0:1 



0:5:9 


0:1:2 
0:0:1 






0:2:2 


0:0:2 


0:1:0 
0:1:0 


0:1:0 






0:3:4 


0:115:7 


1:0:0 


0:1:2 


0:11:14 


0:1:2 


6:2:0 


13:10:2 


1:1:0 


7:1:0 


1:24:13 


0:15:24 


0:15:10 


0:10:16 


0:3:1 


0:2:0 


0:3:2 


4:3:1 






0:2:0 


0:1:2 


0:0:1 


0:9:20 






0:0:2 


0:0:1 










1:1:1 




0:28:79 


0:2:2 
1:0:0 


0:0:2 


0:4:5 


0:2:2 


0:2:1 


0:5:2 
0:2:3 


0:1:0 




0:2:2 
0:0:1 


0:1:2 


0:3:10 




0:1:0 


0:2:4 














0:6:12 




1:24:24 








1:1:0 


2:8:5 


1:1:1 


0:0:1 


0:4:4 


1:24:24 




0:2:1 


















2:3:5 



Table 4 Composition of Acculina-Dulankarella and Parastrophina-Kellerella associations from the Anderken Formation showing number of complete 
shells, ventral and dorsal valves respectively. 







Acculina 


-Dulankarella Association 




Parastrophina- 


-AW/erW/aAssociation 


Sample number 


F-100 


F-1041a 


626 


85258 


8231-40 


628 


8219 


Number of individuals 


168 


113 


112 


32 


11 


40 


2 


Diversity index 


5.07 


1.91 


4.87 


2.60 


1.25 


5.42 




MezotretaP. sp. 


0:1:0 














Schizotreta sp. 






1:0:0 






0:0:1 




Bellimurina sarytumensis 




1:4:0 








0:0:1 




Teratelasmella chugaevae 


12:0:0 




4:0:2 






0:0:1 




Furcitellinae gen. et sp. indet. 


1:1:1 


2:1:2 


1 :0: 1 






0:0:1 




Limbimurinal sp. 


0:0:1 










0:1:1 




Christiania aff. sulcata 












1:6:4 




Christiania egregia 


1:4:1 


2:1:3 


0:2:0 


5:1:0 








Craspedelia tata 


7:2:0 




6:4:0 




0:1:0 


1:0:1 




Acculina kulanketpesica 


12:1:3 


32:3:1 


1:1:1 


4:2:1 


1:5:1 






Dulankarella larga 


25:1:0 


47:0:0 


11:1:0 




1:1:1 






Kajnaria rugosa 


2:0:0 


3:1:0 


1:0:0 






2:1:0 




Mabella conferta 


1:0:0 


3:0:0 








0:1:0 




Shlyginiafragilis 








0:1:0 




0:1:0 




Glxptambonites sp. 


0:2:0 














Sortanella aff. quinquecostata 


1:1:2 




2:3:2 






0:1:1 




Anoptambonites convexus 


12:1:3 




13:6:3 


1:2:1 


1:0:1 






Sowerbyella aff. ampla 


0:2:3 




2:8:0 


0:1:4 








Gacella institata 


3:1:4 




5:1:3 


2:0:0 




0:0:1 




Placotriplesia spissa 


2:1:0 


3:2:1 


1:1:3 


1:0:0 








Triplesia aff. subcarinata 












0:0:2 




Bicuspina rukavishnikovae 


1:0:0 




1:0:0 










Grammoplecia wrighti 






1:0:1 






0:0:2 




Skenidioides sp. 
















Dolerorthis pristina 


1:4:2 




0:1:0 






0:2:2 




Glyptorthis sp. 






1:0:0 






0:2:1 




Austinella sarybulakensis 








3:1:1 








Plectorthis burultasica 




1:1:0 








1:0:0 




Phragmorthis conciliata 






0:1:0 










Parastrophina iliana 


5:1:1 


4:0:0 


7:0:0 


2:0:0 




2:0:1 




Parastrophina plena 


7:0:0 












1:0:0 


Liostrophia pravula 


3:0:2 










1:0:4 




Plectosyntrophia ? unicostata 


2:0:0 




4:0:0 










Schizostrophina margarita 


1:3:0 




2:0:0 






1:0:0 


1:0:0 


Rhxnchotrema akchokense 


2:0:0 




1:0:0 






1:0:0 




Pectenospira pectenata 






1:1:1 










Kellerella misiusi 


3:0:0 




1:0:0 










Nikolaispira guttula 


2:0:0 















UPPER ORDOVICIAN BRACHIOPODS FROM KAZAKHSTAN 



25 



tecturmasi, Dulanaspis levis anderkensis and Styginella 
macrophtalma, whereas Bmnteopsis extraordinaris, Pliomerina sp. 
and Remopleurides sp. are relatively rare. Bivalved molluscs include 
Anderkenia ledomorpha, Clionichia crispa, Edmondia obliqua and 
Praemyopltorisl antiqua (Khalfin 1958), and gastropods are repres- 
ented mostly by Lophospira cribrosa Vostokova. Other fossils include 
unidentified fenestrate and ramose bryozoans of at least three differ- 
ent species, cystoid and crinoid columnals identified by Stukalina 
(1988) as Clivosisystis clivosus, Communicystis communis, 
Digiticrinus levis , Ordinacrinus punctatus, Ristnacrinus bifidus and 
Shizocrinus lentiformis, the cystoidean Polycosmitesl sp. and the 
starfish Stenaster obtusus. 

4. Acculina-Dulankarella Association (Diversity Index 3.14; 
observed range 1.25-5.07, N=5) is characteristic of a nodular algal 
limestone with abundant dasyclad algae which was deposited in the 
base and flanks of carbonate build-ups in the upper Anderken Forma- 
tion. It is a medium to high diversity association defined by the 
occurrence of the plectambonitoideans/\ccM//'«fl kulanketpesica and 
Dulankarella larga. Other brachiopods include Teratelasmella 
chugaevae, Kajnaria rugosa, Gacella institata and Austinella 
sarybulakensis which do not exceed 5% in other associations. Other 
components of the assemblage are taxa common in the Parastrophina- 
Kellerella Association (Tables 4-5), whereas Christiania egregia is 
the only abundant species characteristic also of the Mabella- 
Sowerbyella Association. Mabella conferta and Shlyginia fragilis 
occur sporadically and do not exceed 5% in a particular sample. There 



are abundant green algae: Apidium parvulum, Cyclocrinites nikitini, 
Mastopora reticulata, Mastopora nana and Sinuatipora bucera and 
the red alga Contexta binaria (Gnilovskaya in Nikitin et al. 1974). 
Small organic build-ups up to 30cm across of the algae Girvanella and 
Renalcis are also characteristic and encrust brachiopod shells and 
green algae. The substrate was mainly of silt and lime mud with 
patches of hardground and numerous bioclasts.The abundance of 
brachiopods preserved as conjoined valves (45-64%) in combination 
with the abundant flora of algae suggests quiet environments within 
the euphotic zone of BA-3. Strophomenides (61-90%) constitute the 
most diverse and abundant component of the brachiopod fauna. The 
second most abundant group is the camarelloideans (4-14%), mostly 
Parastwphina iliana and relatively rare Parastrophina plena, Eoana- 
strophia unicostata, Liostrophiapravula and Schizotretina margarita, 
which do not exceed 5% in a particular sample; and orthides, 
rhynchonellides and spire-bearing groups form an insignificant part 
of the association. Among other groups trilobites are the most abun- 
dant. These were partly studied by Weber (1948) and Chugaeva 
(1958), but details remain inadequate. The most characteristic taxa 
are: Illaenus sp.,Acrolichas punctata, Cheirurus sp., Eokosovopeltis 
romanovskyi, Mesotaphraspis spinosus, Pliomerina sulcifrons and 
Sphaerexochus sp. A diverse echinoderm fauna was identified mostly 
from cystoid and crinoid columnals (Stukalina 1988). Among mol- 
luscs the most characteristic is the cephalopod Discoceras 
kazakhstanensisBarskov. The rare coralsAmsassia sp. andLichenaria 
sp., clathrodictyid stromatoporoids, gastropods and bivalved mol- 
luscs are also reported, but remain poorly known. 



Table 5 Composition of Parastrophina- Kellerella Association from the Anderken Formation showing number of complete shells, ventral and dorsal valves 
respectively. 



Sample number 


948 


2538 


8214 


8226 


8217 


8223 


8223a 


8223b 


8256 


Number of individuals 


200 


200 


33 


12 


13 


15 


18 


9 


31 


Diversity index 


2.64 


4.91 


2.63 


1.61 


2.34 


2.58 


3.11 


2.76 


2.33 


Nushbiella dubia 




0:1:1 










0:1:0 






Bellimurina sp. 


0:2:1 


0:1:0 


1:2:0 












0:1:0 


Limbimurina sp. 




0:0:1 












0:0:2 




Christiania aff. sulcata 




0:2:1 


1:4:1 


0:1:0 


0:1:0 


0:2:0 


0:1:1 


0:2:1 




Foliomena prisca 












0:0:1 








Craspedelia tata 


2:2:1 


8:4:3 


1:2:0 








0:1:0 


0:1:0 




Kajnaria rugosa 












1:1:0 








Sortanella aff. quinquecostata 


3:2:1 


4:0:2 
















Anoptambonites convexus 




9:6:2 








0:3:1 






1:3:4 


Sowerbyella aff. ampla 


2:2:1 


3:1:1 


2:2:2 




0:2:0 




3:1:0 


1:0:0 




Trip/esia aff. subcarinata 




2:0:0 
















Placotriplesia spissa 




3:1:7 





2:1 






1:1:0 






0:0:1 


Grammoplecia wrighti 









2:1 














Skenidioides sp. 






1 


0:0 


0:1:0 








0:1:0 




Dolerorthis pristina 


4:5:7 


0:1:1 


2 


1:3 




0:0:1 


0:0:1 




0:1:0 




Glyptorthis sp. 


0:2:1 


2:2:1 


3 


1:1 








0:1:1 




0:0:1 


Plectorthis burultasica 




3:1:1 
















Phaceloorthis sp. 




1:0:0 
















Bowanorthisl devexa 


7:0:0 


5:0:0 
















Phragmorthis conciliata 




2:0:1 










0:0:1 






Parastrophina iliana 


15:0:0 


15:0:1 














2:0:0 


Parastrophina plena 


62:0:0 


17:0:1 


3:0:0 


2:1:1 


2:0:0 




1:0:0 


1:0:0 


5:0:0 


Uistrophina tesikensis 


14:0:0 


















Liostrophia pravula 


7:0:0 


13:0:1 








3:0:0 






3:0:0 


Plectosyntrophia ? unicostata 




1:0:0 






0:0:1 










Sch izostroph ina marga rita 




16:0:0 






0:0:1 


0:0:1 






5:3:2 


Didymelasma cf. transversa 




1:0:0 
















Rhynchotrema akchokense 


2:0:0 


2:0:0 
















Pectenospira pectenata 


14:1:1 


3:0:0 


1:0:0 








3:0:0 






Kellerella misiusi 


15:0:1 


39:0:0 




2:0:0 


5:0:0 




2:0:0 




5:0:0 


Nikolaispira guttula 


13:0:0 


20:0:0 




1:0:0 






2:0:0 






Acculina sp. 




0:1:1 

















26 



L.E. POPOV, L.R.M. COCKS AND I.F. NIKITIN 



5. Parastrophina-Kellerella Association (Diversity Index 3.17, 
observed range 1.61-5.42, N=10) is closely associated with carbon- 
ate build-ups and also belongs to BA-3. These build-ups were 
interpreted by Nikitin et al. (1974) as a chain of bioherms with a 
frame built by the cyanobacterians Girvanella and Renalcis: how- 
ever, micritic limestone usually comprises the most significant part 
of the volume of the rock in the core of a build-up. According to 
Nikitin et al. (1974), these build-ups form a low ridge, raised about 
1.5-3 m above surrounding areas with fine clastic sedimentation. 
Fossils are usually concentrated in pockets of bioclastic limestone 
between individual bioherms and mounds (Samples 948, 2538, 
8223a, 8256). Composition of this association is essentially similar 
to the Acculina-Dulankarella Association, with more than 80% of 
recorded species in common. However, the abundance of 
camarelloideans increases up to 49% (Sample 948) and the archaic 
spire-bearing brachiopods Pectenospira pectenata, Kellerella misiusi 
and Nikolaispira guttula constitute a significant part of the associa- 
tion (21—42% in the most representative samples), whereas in the 
Acculina-Dulankarella Association they are less than 2% (Table 5). 
The relative abundance of strophomenides decreases significantly 
and such genera as Acculina, Dulankarella, Mabella and Shlyginia 
disappear completely. Christiania is represented by the species C. 
aff. sulcata, which is closely linked to this association. Diminution in 
the sizes of the strophomenides might reflect the predominance of 
hard grounds. Taxonomic composition of the association is modified 
in the bedded bioclastic limestone which has large ooids (up to 1 cm 
across) on the top and flanks of the core (Samples 8214, 8217, 8223, 
8223b). Brachiopods are relatively rare and dispersed through the 
rock. Relative abundance of strophomenides, and especially 
Christiania aff. sulcata, increases, whereas spire-bearing brachiopods 
become rare or completely disappear in some samples (8214, 8223, 
9223b). According to the Principal Component Analysis these oc- 
cupy an intermediate position between the cluster formed by samples 
of the Acculina-Dulankarella Association and samples 948, 2538, 
8256, which represent the fully developed Parastrophina-Kellerella 
Association (Fig. 8B). 

The associated trilobite fauna is only partly known and includes 
such taxa as Illaenus sp., Acrolichas punctata, Amphilichas 
punctata, Eokosovopeltis romanovskyi, Glaphurina weberi, 
Mesotaphraspis spinosus, Pliomerina sulcifrons, Selenoharpes sp. 
and Sphaerexochus aff. hisingeri (Weber 1948, Chugaeva 1958, 
Kolobova & Popov, 1986). Crinoid columnals usually represent the 
most important source of bioclasts in the rock. They mostly belong 
to Webericrinus variabilis, Ordinacrinus ordinaris, Malovicrinus 
depressus, Tatjanicrinus crusciformis, Flexicrinus flexus, 
Communicrinus communis and Multifidocrinus mulrifidus ( Stukalina 
1988). 

In the eastern part of the Anderkenen-Akchoku section (Sample 
8226), isolated carbonate build-ups up to 16 m thick appear within 
Unit 2, which is mostly cross-bedded sandstone containing lingulide 
and mollusc associations. It is likely that these build-ups formed 
almost intertidally, but, except for a much lower diversity, the 
brachiopod assemblage retains a relative abundance of spire-bearers 
(Kellerella misiusi) and camerelloideans (Parastrophina p/e/ia) typi- 
cal of the Parastrophina-Kellerella Association, whereas trilobites 
such as Acrolichas sp., Eokosovopeltis romanovskyi , Sphaerexochus 
aff. hisingeri and Illaenus sp. also show close similarity to the 
assemblage from the carbonate unit in the upper part of the Anderken 
Formation. 

6. Zhilgyzambonites-Foliomena Association (Diversity Index 1.21; 
observed range 1.14-130, N=3) is known from three samples col- 
lected from the unit of mudstones and siltstones in the uppermost 



Table 6 Composition of Zhilgyzambonites-Foliomena Association from 
the Anderken Formation showing number of complete shells, ventral and 
dorsal valves respectively. 



Sample number 


2531 


8251 


8255 


Number of specimens 


4 


4 


15 


Diversity index 


1.44 


0.72 


1.48 


Foliomena prisca 


1:0:0 


0:1:0 


1:3:3 


Olgambonites insolita 






0:3:2 


Zhilgyzambonites extenuata 


0:2:1 


0:3:3 


2:5:5 


Kassinella? sp. 






0:0:1 


Chonetoidea sp. 






1:0:0 


Anisopleurella sp. 


0:1:1 







Anderken Formation in the Anderkenyn-Akchoku section (Figs 3, 5, 
Unit 6, Table 6). Brachiopods are a minor part of a trilobite-domi- 
nated benthic assemblage, which includes Amphitrion cf. radians, 
Ampixinella sp., Birmanites almatiensis, Bronteopsis extraordinaris, 
Cheraurus kassini, Cybele weberi, Dionide kazachstanica, 
Dindymene sp., Hamtnatocnemis sp., Microparia speciosa, Ovalo- 
cephalus sp., Granulatagnostus granulatus and Sphaerognostus sp. 
(Chugaeva 1958, Nikitin et al. 1974). Co-occurrence of agnostids, 
cyclopygids and Ovalocephalus allows us to refer this assemblage to 
the Ovalocephalus fauna of Fortey (1997) which is characteristic of 
outer shelf trilobite biofacies corresponding to BA 4-5. Co- 
occurrence of the Foliomena and the Ovalocephalus faunas is 
common in late Caradoc to early Ashgill deep water benthic 
communities in South China (Rong et al. 1994). The Zhilgyzam- 
bonites-Foliomena Association includes only six strophomenide 
genera. Two of them (Olgambonites and Zhilgyzambonites) are 
Kazakhstan endemics, whereas Anisopleurella and Kassinella are 
characteristic of early Foliomena faunas in the Caradoc of China 
(Rong et al. 1999) and elsewhere. 



OVERALL PALAEQECQLOGY 

The sequence of brachiopod associations in the Anderken Formation 
shows an onshore-offshore pattern with a monotaxic lingulide com- 
munity inhabiting mobile sand nearshore, low diversity mollusc and 
brachiopod associations of BA-2 on a shallow clastic shelf, medium 
to high diversity faunas of BA-3 linked with carbonate build-ups, 
and a deep water Foliomena fauna as part of a trilobite-dominated 
benthic assemblage in BA 4-5. In terms of abundance, diversity and 
taxonomic composition, the five associations formed by 
rhynchonelliformean brachiopods can be subdivided into three 
groups, which show little interaction and apparently evolved inde- 
pendently. They are: 1, low-diversity strophomenide-dominated 
Tesikella and Mabella-Sowerbyella Associations of shallow clastic 
shelf corresponding to BA-2; 2, medium to high diversity Acculina- 
Dulankarella and Parastrophina-Kellerella Associations closely 
linked to carbonate build-ups; and 3, a deeper-shelf 
Zhilgyzambonites-Foliomena Association representing an early 
Foliomena Fauna. 

The predominance of strophomenides in environments corre- 
sponding to BA-2, which is usually dominated by rhynchonellides 
and spire-bearing taxa in the late Ordovician and Silurian (Boucot 
1975; Ziegler et al. 1968), is a distinctive feature of the Anderken 
sequence of communities. The Tesikella and Mabella-Sowerbyella 
Associations share eight brachiopod species but Tesikella necopina 
(the index species of the former association) does not usually co- 
occur with Mabella conferta and Shlyginia fragilis. The Tesikella 
necopina Association also demonstrates significant variations in 



UPPER ORDOVICIAN BRACHIOPODS FROM KAZAKHSTAN 



27 



Table 7 Additional list of brachiopod taxa from localities in the Anderken Formation not referred to a particular association 



Sample number 
Number of specimens 



42 
3 



1628 
4 



8215 
10 



620 
13 



8251a 



8135 



Glyptomena onerosa 
Christiania aff. sulcata 
Craspedelia tata 
Dulankarella larga 
Mabella conferta 
Sh lyg in ia frag His 
Anaptambonites orientalis 
Sowerbyella! aff. ampla 
Sowerbyella rukavishnikovae 
Triplesia aff. subcarinata 
Placolriplesia spissa 
Grammoplecia wrighti 
Plectorthis burultasica 
Eodalmanella extera 
Dolerorthis pristina 
Pionodema opima 



0:1:0 



1:0:0 
0:0:1 



2:0:0 



2:0:0 



1:3:0 



1:4:1 
0:1:0 



2:1:0 
1:0:0 



0:1:0 
5:7:2 



2:1:8 
1:0:1 
2:0:1 



0: 1 :0 



0:1:0 



2:0:0 



relative abundance and taxonomic composition from one sample to 
another, which may reflect its opportunistic character and higher 
environmental stress, whereas the Mabella-Sowerbyella Associa- 
tion, in spite of its low diversity (4 to 8 genera per sample), shows a 
relatively constant taxonomic composition in most samples (Table 
3), and this association forms a compact cluster in the Principal 
Components Analysis (Fig. 9). Comparison with somewhat younger 
Ctenodonta-Sowerbyella,Altaethyrella-Nalivkinia (Pronalivkinia) 
and Dinorthis Associations of the lower Dulankara Formation which 
inhabited similar environments on the shallow clastic shelf on the 
Chu-Ili Plate in the late Caradoc-early Ashgill (Popov et al. 2000), 
suggest a rapid faunal turnover. In these younger faunas Sowerbyella 
and Shlyginia retained their dominant position and Anoptambonites 
also remained a characteristic minor component, but most of the 
Anderken genera disappeared (e.g. Tesikella, Eodalmanella and 
Pionodema) or moved into the middle shelf {Mabella and 
Glyptomena) and were replaced by rhynchonellides such as 
Altaethyrella and atrypides such as Nalivkinia (Pronalivkinia). In 
contrast, the orthides (e.g. Plaesiomys, Bokotorthis and Dinorthis) 
became increasingly abundant. 

The diverse Acculina-Dulankarella and Parastrophina-Kellerella 
Associations, which were closely linked with carbonate build-ups, 
have little in common with the faunas which inhabited shallow 
clastic shelves nearby. In the Acculina-Dulankarella Association 
strophomenides remain the most abundant brachiopods but they are 
mostly different genera, such as Bellimurina, Teretelasmella, 
Craspedelia, Acculina, Dulankarella. Kajnaria and Sortanella. 
Anoptambonites occurs in both associations, but as different species. 
Mabella and Shlyginia are mostly confined to the Mabella- 
Sowerbyella Association, as well as occurring in a few samples of the 
Acculina-Dulankarella Association, but as less than 5% of the 
sample (Table 4). They disappear in the Parastrophina-Kellerella 
Association (Table 5), which is possibly the earliest known 
brachiopod assemblage in which pentamerides together with spire- 
bearing taxa come to a dominant position. In contrast to the 
brachiopod fauna of the shallow clastic shelf, assemblages associ- 
ated with carbonate build-ups did not undergo any significant 
taxonomic change during the Caradoc. The late Caradoc to early 
Ashgill brachiopod fauna of the Dulankara carbonate mud-mound in 
the north Betpak-Dala Desert (Nikitin et al. 1 996), which was also on 
the same Chu-Ili Plate, retained a close similarity to the Anderken 
fauna and contained 14 genera in common including Parastrophina 
and the early athyridides Kellerella and Nikolaispira. The 



strophomenide component was largely unchanged and such genera 
as Bellimurina, Limbimurina, Christiania, Craspedelia, Sortanella 
and Anoptambonites are common to both faunas. 

The significance of the Foliomena fauna was discussed by Cocks 
& Rong (1988) and Rong et al. (1994, 1 999). In the Lower to Middle 
Caradoc it was confined mostly to South China and only in the 
Ashgill did it become cosmopolitan. In Kazakhstan there are no 
previous reports of the occurrence of Foliomena, but the associated 
Ovalocephalus trilobite fauna occurs in many outer shelf environ- 
ments from the Middle Caradoc (Apollonov 1975; Nikitin et al. 
1974). In the Anderken Formation Foliomena itself is not restricted 
to BA 4—5, but occurs occasionally in the Parastrophina-Kellerella 
(Sample 8223) and Mabella-Sowerbyella (Sample 8228) Associa- 
tions. However, the other taxa, which include the two new genera and 
species Olgambonites insolita and Zhilgyzambonites extenuata 
together with rare Anisopleurella, Chonetoidea and Kassinella, are 
not present in shallow shelf associations. In addition, there are seven 
localities (Table 7) whose brachiopods cannot be referred with 
confidence to any of the six named associations. 

Although comprehensive comparisons of the Anderken faunas 
with other contemporary brachiopods from elsewhere are beyond 
the scope of this paper, it is worth noting here that the total Anderken 
assemblage has much in common with that described from north- 
west China (Fu 1982). 



SYSTEMATIC PALAEONTOLOGY 

Figured and cited specimens are housed in the Natural History 
Museum. London (BB and BC collection numbers). Institute of 
Geological Sciences, Almaty, Kazakhstan (IGNA collection num- 
bers), and in the CNIGR Museum, St. Petersburg, Russia (CNIGR 
collection numbers). All the quoted sample numbers are from the 
Anderken Formation except where stated. Bibliographical refer- 
ences to families and above are omitted if they are in the Treatise on 
Invertebrate Paleontology (Kaesler 2000). 

Abbreviations given in tables of measurements and in the text are: 
Lv, Ld - sagittal ventral and dorsal valve length; W - maximum 
width; T - maximum thickness; Ml, Mw - length and width of the 
muscle field; Sw, St - width and height of tongue in the ventral valve; 
BB1, BBw length and distance between outer margins of 
brachiophores or socket ridges; SI - length of median ridge; LP1, 



28 

LPw length and width of lophophore platform; X - mean; S - 
standard deviation from the mean; r- coefficient of correlation; OR 

- observed range; max. - maximum value; min. - minimum value; N 

- number of measured or counted specimens. 



L.E. POPOV, L.R.M. COCKS AND I.F. NIK1TIN 

species, which makes precise comparison difficult. The main differ- 
ence of the Kazakh species from the latter is the less elongate shell 
outline, which is no more than twice as long as wide. A detailed 
description, discussion and the basic statistics of this species was 
provided by Popov ( 1980). 






Order LINGULIDA Waagen, 1885 
Superfamily LINGULOIDEA Menke, 1828 

Family OBOLIDAE King, 1846 

Subfamily GLOSSELLINAE Cooper, 1956 

Genus ECTENOGLOSSA Sinclair, 1945 

Type species. Lingula lesueueri Rouault, 1850, from the Arenig 
of Normandy, France. 

Ectenoglossa sorbulakensis Popov, 1980 PI. 1, figs 1^1 

1980 Ectenoglossa sorbulakensis Popov: 142, pi. 1, figs 1-4. 
1984 Ectenoglossa sorbulakensis Popov; Nikitin & Popov in 
Kleninaera/.: 142, pi. l,figs 1^1. 

Holotype. CNIGR 1/11523, ventral internal mould, from the 
Anderken Formation, locality 1024, east side of Karatal valley. 

Material. Six pairs of conjoined valves, 26 ventral and 3 1 dorsal 
valves from Sample 8130a, Anderkenyn-Akchoku; Samples 8227- 
10, 8227^40 (BC 57370-73), 8227-80, Buldukbai Akchoku; 8228a, 
east side of Kopalysai; all Chu-Ili Range; Sample 1024, east side of 
Karatal Valley, south Betpak-Dala. 

Description. Shell equivalved, elongate, subrectangular in out- 
line, about 1 90% as long as wide with maximum width at mid-length, 
ornamented by fine concentric fila about 13-15 per mm. Ventral 
valve very gently convex with narrow, triangular pseudointerarea, 
mainly occupied by deep pedicle groove and small propareas crossed 
by flexure lines. Dorsal valve gently convex, lacking pseudointerarea. 
Ventral interior with pair of slightly diverging, elongate, suboval 
umbonal muscle scars flanked laterally by pair of short diverging 
ridges; pedicle nerve impression well defined. Dorsal interior with 
weak median ridge. 

Discussion. Ectenoglossa sorbulakensis closely resembles 
Ectenoglossa minor Zhan & Cocks (1998: 14) from the Lower 
Ashgill of South China in having a strongly elongate equivalved 
shell with subparallel lateral margins, rudimentary ventral 
pseudointerarea and propareas with well defined flexure lines. How- 
ever, the interiors of both valves are weakly impressed in the Chinese 



Superfamily DISCINOIDEA Gray. 1840 

Family TREMATIDAE Schuchert, 1893 

Genus TREMATIS Sharpe, 1848 

TYPE SPECIES. Orbicula terminalis Emmons, from the Trenton 
Group (Caradoc), New York, U.S.A. 



Trematisl sp 



PI. 1, figs 5, 6 



Material. Figured dorsal valve, BC 57375 (L=14.8, W=14.4) 
and another dorsal valve, Samples 127, 8228, Kopalysai. 

DESCRIPTION. Shell subcircular, ornamented by radial capillae of 
about 5 per 1 mm enlarged in number by intercalation and with 
radially arranged rounded or transversely suboval pits in the 
interspaces between capillae. Ventral valve unknown. Dorsal valve 
gently and unevenly convex in transverse profile with maximum 
height about one-third of the valve length from the marginal umbo. 
Dorsal interior unknown. 

Discussion. These specimens are similar to the shells described 
by Cooper (1956: 275) as Trematis sp. 3 from the Cyrtonotella Zone 
of the Edinburg Formation (Caradoc) of Virginia in their pits and 
radial ornament. The ventral valve in both unnamed species remains 
unknown, and therefore the generic attribution is provisional. 



Family DISCINIDAE Gray, 1840 

Subfamily ORBICULOIDEINAE Schuchert, 1929 

Genus SCHIZOTRETA Kutorga, 1848 

Type SPECIES. Orbicula elliptica Kutorga, presumably Volkhov or 
Kunda Stage (Upper Arenig-Lower Llanvirn), vicinity of St. 
Petersburg. Russia. 



Schizotreta sp. 



PI. 1, figs 7-10 



Material. One pair of conjoined valves, one ventral and two 
dorsal valves from Samples 100 (BC 57590) and 626, Anderkenyn- 
Akchoku section; Samples 628 (BC 56825), 2538, Kujandysai section 



PLATE 1 

Figs 1-4 Ectenoglossa sorbulakensis Popov. Sample 8227—40, Buldukbai-Akchoku section, west side of Kopalysai. 1, BC 57370, ventral exterior, x 2. 2, 

BC 57372, dorsal internal mould, x 2. 3, BC 57371, dorsal exterior, x 2. 4, BC57373, ventral internal mould, x 2. 
Figs 5, 6 Trematis ? sp. Sample 8228, east side of Kopalysai, BC 57375, dorsal exterior and surface ornament, x 6, x 2. 

Figs 7-10 Schizotreta sp. Sample 100, Anderkenyn-Akchoku section, BC 57590, conjoined valves, posterior, dorsal, lateral and ventral views, x 2. 
Fig. 11, 12 MesotretaP. sp. Sample 100, Akchoku Mountain, Anderkenyn-Akchoku section, CNIGR 1/12361, ventral exterior and lateral view, x 8. 
Figs 13, 14 Nushbiella dubia Popov. Sample 2538, Akchoku Mountain, Kujandysai section. BC 57591 ; 13, dorsal exterior, x 8; 14, ventral exterior, x 8. 
Fig. 15 Paracraniops sp. Sample 8128a, Anderkenyn-Akchoku section, BC 57377, dorsal internal mould, x 5. 
Figs 16-21 Longvillia lanx (Popov), Sample 1018, area 7 km southwest of Karpkuduk well, Kotnak mountains, south Betpak-Dala. 16, 17. CNIGR 27/ 

1 1989, latex cast of cardinalia and dorsal interior, x 5, x 2. 18-20, CNIGR 27/11989, dorsal, ventral and lateral views of conjoined valves, x 1. 21, 

CNIGR 30/1 1989, holotype, ventral internal mould, x 2. 
Figs 22-28 Bellimurina (Bellimurina) sarytumensis sp. nov. 22, 23, Sample 8214, BC 57379, holotype, Anderkenyn-Akchoku section, ventral exterior 

and lateral view, x 2. 24, 25. Sample 2538, Akchoku Mountain. Kujandysai section, BC 57378, conjoined valves, dorsal and ventral views, x 3. 26, 

Sample 1041a, Burultas Valley, BC 57364, dorsal internal mould, x 2. 27, Sample 100, Anderkenyn-Akchoku section, BC 57380, ventral exterior, x 3. 28, 

Sample 2538, CNIGR 10/12361, dorsal external mould, x 3. 
Fig. 29 Dolerorthis expressa Popov. Sample 1018, 7 km southwest of Karpkuduk well, Kotnak Mountains, BC 57368, ventral internal mould, x 1.5. 
Figs 30, 31 Furcitellinae gen. et sp. indet. Sample 628, west side of Kujandysai, BC 57381, ventral and dorsal views of conjoined valves, x 1.5. 



UPPER ORDOVICIAN BRACHIOPODS FROM KAZAKHSTAN 



29 




30 



L.E. POPOV, L.R.M. COCKS AND I.F. NIKITIN 



Description. Shell planoconvex, subcircular, ornamented by 
strong rounded irregular concentric rugellae of 10-11 per 3 mm. 
Ventral valve low, subconical with maximum height at the umbo 
which is situated about 10% of the valve length from the posterior 
margin. Pedicle foramen small, rounded, located at the end of narrow 
pedicle track covered by listrium. Dorsal valve flat with submarginal 
umbo. Interior of both valves not observed. 



Nushbiella dubia (Popov, 1977) 



PL 1 , figs 13, 14 



Measurements. 
T=4.8. 



BC 57590, conjoined valves, L=14.8, W=14.4, 



Discussion. These specimens resemble Schizotreta triangularis 
Popov (Nikitin et al. 1996: 85) from the late Caradoc Dulankara 
Regional Stage of the northern Betpak-Dala Desert, Central 
Kazakhstan, but differ in their subcircular valve outline, ventral 
umbo situated close to the posterior margin and strong, irregular 
concentric rugellae. In concentric ornament they somewhat resemble 
Schizotreta corrugata Cooper ( 1956: 277) from the Llandeilo Pratt 
Ferry Formation of Alabama, USA, but can be distinguished in 
having a larger size, more circular shell outline and more densely 
spaced concentric rugellae. 

Order SIPHONOTRETIDA Kuhn, 1949 

Superfamily SIPHONOTRETOIDEA Kutorga, 1848 

Family SIPHONOTRETIDAE Kutorga, 1848 

Genus MESOTRETA Kutorga, 1848 

Type species. Siphonotreta tentorium Kutorga, 1848: 270, from 
the Arenig (Volkhov Regional Stage), north-western Russia. 

DISCUSSION. The type material of Mesotreta tentorium is lost and 
its taxonomic position within the Siphonotretoidea is unclear. At 
present, our knowledge of the type species of this genus is based on 
a single incomplete ventral valve from the Upper Volkhov Stage 
(Holmer & Popov 2000: Fig. 79,2). Mesotreta is unique among the 
Siphonotretida because of its low conical ventral valve with small, 
slightly eccentric pedicle foramen, but it possesses the hollow spines 
characteristic of this order. The ventral interior and dorsal valve of 
Mesotreta remain unknown. 



Mesotreta? sp. 



PI. l,figs 11, 12 



1 986 Nushbiella dubia (Popov); Kolobova & Popov: pi. 1 , fig. 1, 
non fig. 2. 

Material. One ventral valve (CNIGR 1/12361) from Sample 
100, Anderkenyn-Akchoku section. 

DESCRIPTION. Valve slightly transverse, elliptical in outline; low, 
subconical with eccentric umbo at 20% of valve length from the 
posterior margin. Foramen small, slightly elongate, suboval. Con- 
centric ornament of up to 16 thin, evenly spaced, crowded growth 
lamellae and numerous fine hollow spines. Ventral interior and 
dorsal valve unknown. 

DISCUSSION. This unnamed species resembles Siphonotreta tento- 
rium in having a low conical ventral valve lacking a well-defined 
pseudointerarea and small umbonal pedicle foramen, but the 
Kazakhstanian specimen differs in its strongly developed lamellose 
ornament and more posterior ventral umbo. 



Genus NUSHBIELLA Popov in Kolobova & Popov, 1986 

TYPE SPECIES. Multispinula? dubia Popov, 1977; from the 
Bestamak Formation, Tselinograd Regional Stage (Llandeilo), 
Chingiz Range, Kazakhstan. 



1977 Multispinula? dubia Popov: 104, pi. 25, figs 8-11. 

1986 Nushbiella dubia (Popov) Kolobova & Popov: 251, pi. 1, 

fig. 2, non fig. 1 . 
2000 Nushbiella dubia (Popov) Holmer & Popov: fig. 79, 6a, b. 

Holotype. CNIGR 1/10847, ventral valve, Bestamak Formation 
(Llandeilo), locality 553a, east side of Chagan River near Konur- 
Aulie cave. Chingiz Range, Kazakhstan. 

Material. One ventral and one dorsal valve from Sample 8223a, 
Anderkenyn-Akchoku section; Sample 2538 (BC 57591), Kujandysai 
section. 

Discussion. The specimens from the Anderken Formation have 
no significant differences from the rather older topotypes in orna- 
ment and external shell morphology. 



Order CRANIOPSIDA Gorjansky & Popov, 1985 

Superfamily CRANIOPSOIDEA Williams, 1963 

Genus PARACRANIOPS Williams, 1963 

TYPE SPECIES. Craniops pararia Williams, 1962, from the Lower 
Ardmillan Series (Caradoc), Girvan, Scotland. 



Paracraniops sp. 



PI. 1, fig. 15 



Material. One dorsal internal mould, BC 57377, from Sample 
8128a, Anderkenyn-Akchoku section. 

Discussion. A single specimen represents the earliest record of 
craniopsides from Kazakhstan. There is little doubt of the generic 
attribution, but there is insufficient material to allow detailed com- 
parison with other named species of the genus. 



Order STROPHOMENIDA Opik, 1934 

Superfamily STROPHOMENOIDEA King, 1846 

Family STROPHOMENIDAE King, 1846 

Subfamily STROPHOMENINAE King, 1846 

Genus LONGVILLIA Bancroft, 1933 

Type SPECIES. Orthis grandis J.de C. Sowerby, 1839, from the 
Cheney Longville Flags (Lower Caradoc), Shropshire, England. 



Longvillia lanx (Popov, 1985) 



PI. 1, figs 16-21 



1985 Strophomena /<mv Popov: 58; pi. 1, fig. 13; pi. 2, figs 12, 13; 

pi. 3, fig. 1. 
1 985 Strophomena digna Popov: 67 nomen erratum, pi. 1 , fig. 13. 

Holotype. CNIGR 30/11989, ventral internal mould, from the 
Anderken Formation, Sample 1 1 8a, 7 km south-west of Karpkuduk 
well, Kotnak Mountains, refigured here (PI. 1, fig. 21). 

Material. Two pairs of conjoined valves, 1 1 ventral and 8 dorsal 
valves from Samples 1018, 1018a, 7 km south-west of Karpkuduk 
well, Kotnak Mountains. 

Description. Shell very gently convexiconcave, slightly trans- 
verse, subrectangular in outline with maximum width about 
one-quarter of the valve length from the hinge line. Anterior com- 
missure rectimarginate. Ventral valve gently and evenly concave in 
profile with a slightly raised, pointed umbo and steeply apsacline 
ventral interarea bearing large, convex pseudodeltidium. Dorsal 
valve evenly convex with flattened umbonal area. Radial ornament 



UPPER ORDOVICIAN BRACHIOPODS FROM KAZAKHSTAN 



31 



unequally parvicostellate in posterior half of mature valves with 2- 
4 parvicostellae between accentuated ribs, becoming near equally 
parvicostellate anteriorly with 5-8 ribs along the anterior margin. 
Ventral interior with strong teeth and low, diverging dental plates 
continuing anteriorly as straight muscle bounding ridges bordering 
posteriorly the open subtriangular muscle field which is about 30% 
as long as the valve. Adductor scars narrow, weakly impressed, 
flanked laterally by slightly longer diductor scars. Dorsal interior 
with cardinal process, low, widely diverging, oblique socket ridges 
and a short median ridge up to 40% valve length. Adductor field 
weakly impressed. 

Measurements. CNIGR 29/11989, conjoined valves, L=22.5, 
W=26.4, Iw=23.6, T=5. 1 ; CNIGR 30/1 1989, ventral internal mould, 
holotype, L=26.8, W=28.3, Ml=8.8, Mw=9.6; CNIGR 35/11989, 
dorsal internal mould, L=16.7, W=18.5, T=3.5, Sl=7.8. 

Discussion. This species was originally referred to Strophomena, 
but it is characterised by an open ventral muscle field bounded 
posterolaterally by dental plates and short, diverging bounding ridges, 
a short dorsal median ridge and the absence of side septa, all 
suggesting attribution to Longvillia. It differs from the type species 
Longvillia grandis in its less transverse outline, with the maximum 
width slightly anterior to the hinge line, the radial ornament becom- 
ing nearly regular in the anterior half of the shell, and in possessing 
a somewhat stronger median ridge. 



Subfamily FURCITELLINAE Williams, 1965 
Genus BELLIMURINA (BELLIMURINA) Cooper, 1956 

TYPE SPECIES. Leptaena charlottae Winchell & Schuchert, from 
the Caradoc of Minnesota, U.S.A. 

Bellimurina (Bellimurina) sarytumensis sp.nov. PL 1 , figs. 

22-28. 

1986 Bellimurina rudis [sic] Lu; Kolobova & Popov: pi. 1, fig. 
10. 

ETYMOLOGY. After Sarytuma Well in the Betpak-Dala Desert. 

Holotype. BC 57379, PL 1, figs 22, 23, a ventral valve from 
Sample 8214, Anderkenyn-Akchoku section. 

MATERIAL. Two pairs of conjoined valves, ten ventral and four 
dorsal valves from Samples 100 (BC 56826, BC 57380), 620 (BC 
56827-28) and 626 (BC 56829-3 1 ), Anderkenyn-Akchoku section; 
Sample 8214 (BC 56839-40, BC 57379), west side of Ashchisu 
River; Samples 628 (BC 56837), 2538 (BC 56838, BC 57378) and 
8256 (BC 56841), Kujandysai section; Sample 948 (BC 56832-36), 
Tesik River; Samples 390 (BC 57365), 1041a (BC 57364), Burultas 
Valley. 

Description. Shell concavoconvex, slightly transverse, sub- 
rectangular in outline, about 70% as long as wide with maximum 
width at mid-length with rounded cardinal extremities and 
rectimarginate anterior commissure. Ventral valve convex with maxi- 
mum height at geniculation (about three-quarters valve length). 
Ventral interarea apsacline with well developed pseudodeltidium 
perforated apically by a minute foramen. Dorsal valve flattened with 
geniculation near the anterior margin. Dorsal interarea low, linear, 
anacline with a well-developed chilidium. Radial ornament un- 
equally parvicostellate with 6-7 accentuated ribs originating at the 
umbo and up to 7 strong ribs in interspaces. About 4-7 fine 
parvicostellae per mm along the anterior margin. Concentric ornament 



Table 8 Measurements of ventral valves of Beillimurina sarytumensis sp. 
nov from samples 100, 626 and 8214 from Anderkenyn-Akchoku 
section. Sample 2538 from Kujandysai section and Sample F- 1041a 
from Burultas valley. 



Lv 



W 



LAV 



N 


6 


X 


7.8 


S 


3.62 


MIN 


5.2 


MAX 


14.8 



6 


6 


1.1 


69.4% 


4.35 


6.8 


8 


56.5% 


9.5 


75.9% 



of fine, slightly uneven rugellae about 2 per mm, covering all the 
posterior half of the shell. 

Ventral interior with small teeth supported by low, divergent 
dental plates and small, weakly impressed subtriangular muscle 
field. Dorsal interior with bilobed cardinal process, widely flaring 
socket plates, poorly impressed median septum extending about 
one-third valve length, poorly impressed pair of side septa subparallel 
to the median septum and not extending beyond it. 

Measurements. (435/12375) conjoined valves, L=6.4, W=8.3, 
T=3.0; (436/12375) ventral valve, L=15.4, W=21.3, T=5.4. 

Discussion. These shells are most similar to, and possibly 
conspecific with, the specimens described by Nikitin & Popov 
(1996: 16, figs 6J-N) as Bellimurina? sp. from the Dulankara 
Regional Stage of north Betpak-Dala in general shell shape and 
radial ornament with 6-7 accentuated primary ribs, but the dorsal 
interior in specimens from the Dulankara Stage remains unknown. 
The Kazakh shells are similar to the specimens described as 
Kiaeromena longxianensis Fu, 1982 from the Pinling Formation of 
northwest China in size, ornament and weakly geniculated lateral 
profile, but further comparison is difficult because of poor descrip- 
tion and insufficient illustrations of that species, although it is 
unlikely to belong to Kiaeromena, which is known only from the 
Baltic. Cooper (1956) described eight species of B. (Bellimurina) 
from the Caradoc of Laurentia, of which he named six, but the dorsal 
interiors do not match B. (B.) sarytumensis, nor do those of B. (B.) 
rudis Xu, Rong & Liu (1974) from the early Caradoc Shitzupu 
Formation of South China. B. (B.) quadrata Fu (1982) appears more 
quadrate in outline and its interior is not illustrated (see also under 
Limbimurina? sp. below). 



Genus TERATELASMELLA Laurie, 1991 

Type SPECIES. Teratelasmella plicata Laurie, 1 99 1 , from the Up- 
per Cashions Creek Limestone (Lower Caradoc ), Tasmania, Australia. 



Teratelasmella chugaevae sp. nov. 
10, 11. 



PL 2, figs 10-20; Figs 



ETYMOLOGY. In memory of the late Marina Chugaeva to honour 
her outstanding trilobite work. 

Holotype. BC 57392, PL 2, figs 14-18, conjoined valves, from 
the Anderken Formation, Sample 626, Anderkenyn-Akchoku section. 

Material. 1 6 pairs of conjoined valves and 3 dorsal valves from 
the Anderkenyn-Akchoku section, Samples 100 (BC 56842-5 1 , BC 
57390-91) and 626 (BC 56853-57, 57392); Kujandysai section. 
Samples 628 (BC 56858) and 85258 (BC 56852). 

Description. Shell strongly dorsibiconvex, transverse, suboval, 
about 70% as long as wide, with maximum width at mid-length, and 



32 



L.E. POPOV. L.R.M. COCKS AND I.F. NIKITIN 



1(0.5) 2(1.3) 




16(6.5) 



10 

I 



20 mm 

j i 



Fig. 10 Transverse serial sections of conjoined valves of Teratelasmella chugaevae sp. nov. from Sample 100. Distance in mm is measured from the 
posterior tip of ventral beak. Dorsal valve uppermost. Also lateral view to show section positions and schematic reconstruction. 



90% as thick as long. Anterior commissure strongly uniplicate. 
Ventral valve moderately convex with maximum thickness near 
quarter valve length. Interarea high, triangular, apsacline with 
delthyrium completely covered by pseudodeltidium perforated by a 
small umbonal foramen. Ventral sulcus originating at the umbonal 
area, strongly deepening anteriorly and ending with prominent, 
semielliptical tongue inclined at less than a right angle to the com- 
missural plane. Flanks of the valve flattened, slightly inclined to the 
commissural plane. Dorsal valve very strongly convex, with a low, 
anacline interarea. Strong median fold, rounded in cross-section, 
with steep lateral slopes originating near the beak. Very weak and 
narrow dorsal median sulcus in the umbonal area of some specimens. 
Radial ornament coarsely parvicostellate with 5 ribs per 3 mm along 
the posterior margin of mature specimens. 

Interiors of both valves were studied in transverse sections (Figs 
10, 11). Ventral valve with strong teeth supported by high dental 
plates continuing anteriorly as muscle bounding ridges. Low median 
ridge in the anterior half of the ventral muscle field. Dorsal valve 
interior with bilobed cardinal process and low, curved socket ridges. 
Median septum high, triangular, blade-like, extending anteriorly to 
mid-valve, flanked laterally by a pair of short side septa. Adductor 
field raised anteriorly and bordered by a high rim. 

DISCUSSION. This species is similar only to Teratelasmella plicata 



Laurie, 1991. but it can be distinguished in being larger (up to 25.6 
mm wide), with a strongly dorsibiconvex lateral profile, a deep 
ventral sulcus, a high dorsal median fold originating in the umbonal 
area rather than at the mid-valve as in the type species, and coarser 
radial ornament. 

Furcitellinae gen. et sp. indet. 

PI. 1, figs 30, 31, PI. 2, figs 1-4 

Material. Four pairs of conjoined valves, two ventral and six 
dorsal valves from Samples 100 (BC 56859, 57382, 57384) and 626, 
Anderkenyn-Akchoku section; Sample 628 (BC 56577, 56863, 
57381) and 8220 (BC 56578), Kujandysai section; Sample 1041a 
(BC 56860-62, 57383), Burultas Valley. 

Description. Shell convexoplane, transverse, subrectangular in 
outline with maximum width near mid-length. Cardinal extremities 
nearly right angled. Anterior commissure rectimarginate. Ventral 
valve almost flat with acute and slightly erect beak. Ventral interarea 
low apsacline with convex pseudodeltidium. Dorsal valve evenly 
convex with low, orthocline interarea and well-developed chilidium. 
Radial ornament parvicostellate. Up to 10 mm from the umbo the 
ribs are nearly equal in size and are 7-9 per 3 mm. In larger 
specimens 2-5 finer costellae become inserted between the larger 



UPPER ORDOVICIAN BRACHIOPODS FROM KAZAKHSTAN 



33 




9 (5.0) 



5 10 mm 

_i i i i i i 



Fig. 11 Transverse serial sections of Teratelasmella chugaevae sp. nov. from Sample 100. Distance in mm is measured from the posterior tip of ventral 
beak. Dorsal valve uppermost. Also lateral view to show section positions and schematic reconstruction. 



ribs and ornament becomes unequally parvicostellate, with up to 4 
accentuated ribs and 9-15 parvicostellae per 3 mm along the anterior 
margin. 

Ventral interior with delicate teeth and long, divergent dental 
plates. Ventral muscle field subtriangular, open anteriorly. Dorsal 
interior with bilobed cardinal process. Other characters of dorsal 
interior unknown. 

Discussion. Within the Furcitellidae as revised by Cocks & Rong 
(2000) these shells are comparable with Quondongia (Percival 
1 99 1 : 1 5 1 ) and Molongcola (Percival 1 99 1 : 1 53) from the Caradoc of 



Australia in their planoconvex lateral shell profile and in the absence 
of geniculation, but they differ from Molongcola in possessing 
parvicostellate ornament. The absence of information on the dorsal 
interior makes precise generic attribution impossible. 

Family GLYPTOMENIDAE Cooper, 1956 
Genus GLYPTOMENA Cooper, 1956 

TYPE SPECIES. Glyptomena sculpturata Cooper, 1956. from the 
Chatham Hill Formation (Llandeilo), Virginia, U.S.A. 



34 



L.E. POPOV, L.R.M. COCKS AND I.F. NIKITIN 




UPPER ORDOVICIAN BRACHIOPODS FROM KAZAKHSTAN 

Glyptomena onerosa Popov, 1980 PI. 2, figs 5-9 

1980 Glyptomena onerosa Popov: 152, pi. 2, figs 5-7. 

HOLOTYPE. CNIGR 50/1 1523, Anderken Formation, from Sample 
100b, Anderkenyn-Akchoku section. 

Material. 14 ventral and 24 dorsal valves from Samples 100b, 
843 (BC 57387), 8135, 8137, 8235 (BC 57386), Anderkenyn- 
Akchoku section; Sample 7613 (BC 57388), Kujandysai section; 
Samples 8229 (BC 57385), 8257 (BC 56864-65), Buldukbai- 
Akchoku section; Sample 1018, 7 km SW of Karpkuduk well, 
Kotnak Mountains. 

Description. Shell concavoconvex, semielliptical in outline, about 
60% as long as wide, with maximum width at the hinge line. Cardinal 
extremities slightly acute. Anterior commissure rectimarginate. Ven- 
tral valve moderately convex in lateral profile with maximum 
thickness at about one-third valve length. Ventral interarea low, 
apsacline, with small apical pseudodeltidium. Dorsal valve flattened 
with dorsal geniculation at about 75% of valve length from the 
umbo. Dorsal interarea low, anacline with well developed, broad, 
convex chilidium. Radial ornament inequally parvicostellate with up 
to three generations of accentuated ribs separated by 2-5 
parvicostellae in the interspaces. Number of ribs along the anterior 
margin of mature specimens varying from 11 to 15 per 3 mm. 
Concentric ornament of fine, regularly spaced concentric fila, about 
20-25 per mm. 

Ventral interior with strong teeth bearing rows of crenulations on 
the outer side and long, widely divergent dental plates. Ventral 
muscle field heart-shaped about one-third as long as the valve with 
strongly impressed diductor scar somewhat longer, but not enclosing 
slightly raised, narrow triangular adductor track. Dorsal interior with 
bilobed cardinal process on a low notothyrial platform and widely 
diverging socket ridges subparallel to the hinge line. Sockets deep, 
transverse, bearing strong vertical ridges on the anterior slope. 
Adductor scars weakly impressed, bisected by the fine median 
septum extending anteriorly about 60% of valve length. Pair of inner 
side septa about the same length as the median septum, slightly 
divergent proximally and curved towards the anterior of the median 
ridge near the mid-valve. Outer side septa very fine or completely 
absent in some specimens. 

DISCUSSION. This species is similar to Glyptomena sculpturata 
Cooper, 1956, but differs in having a geniculated dorsal valve and 
more densely accentuated ribs. 



Limbimurina? sp. 



35 
PI. 3, fig. 1 



Material. One ventral and five dorsal valves from Samples 100, 
626 (BC 57395), 8223b (BC 56867-68), Anderkenyn-Akchoku 
section; Samples 628 and 2538 (BC 56866), Kujandysai section. 

Description. Shell flattened, transverse, subrectangular in out- 
line; strongly geniculate ventrally with a trail up to 4 mm long 
inclined at nearly right angles to the commissural plane. Cardinal 
extremities nearly right angled. Ventral valve strongly flattened 
posteriorly to the geniculation. with gently convex umbonal area. 
Ventral interarea low, planar, apsacline with apical pseudodeltidium. 
Dorsal valve with strong angular concentric rugae accentuating the 
geniculation. Dorsal interarea anacline. Radial ornament unequally 
parvicostellate with 4-5 parvicostellae per mm along the anterior 
margin. Concentric ornament of oblique rugellae crossing each other 
at less than 30-40° and covering all the valve surface between hinge 
line and geniculation. Interior of both valves unknown. 

Measurements, dorsal valve, L= 1 4. 1 , W=29.0. 

Discussion. Generic assignment of the Kazakh specimens to 
Limbimurina is based mostly on the distinctive shell shape, with 
strong geniculation enhanced by the characteristic concentric frill 
and the irregular oblique rugellae forming an interference pattern 
with concentric rugae in the posterior half of the shell. The speci- 
mens from the Anderken Formation differ from Limbimurina 
brevilimbata Cooper, 1956 from the Edinburg Formation of Virginia 
and L. insueta Cooper, 1956 from the Rodman Formation of Penn- 
sylvania in the strongly transverse outline of the shell. The trail in the 
Kazakh shells is considerably higher than in L. insueta, but not as 
high as in L. brevilimbata. 

Similar shells were also described as Limbimurina 1 . sp. from the 
Dulankara Regional Stage (Upper Caradoc) of north Betpak-Dala 
(Nikitin & Popov 1996). They differ from the Anderken specimens 
in their more densely spaced parvicostellae, which vary from 1 1 to 
14 per mm along the anterior margin. 

A similar species is Bellimurina quadrata Fu, 1982, from the 
Pinling Formation of Northwest China, which also possesses a 
geniculation as well as a characteristic concentric frill, and irregular 
oblique rugellae forming characteristic interference patterns; but 
precise comparision is difficult because of the short description and 
insufficient illustrations. It most likely belongs to Limbimurina, but 
no detailed information was provided on the dorsal interior (Fu 
1982:122, pi. 35, figs 20-21). 



Family LEPTAENIDAE Hall & Clarke, 1894 
Genus LIMBIMURINA Cooper, 1956 

TYPE SPECIES. Limbimurina insueta Cooper, 1956, from the 
Nealmont Formation (Caradoc), Pennsylvania, U.S.A. 



Family CHRISTIANIIDAE Williams, 1952 
Genus CHRISTIANIA Hall & Clarke, 1892 



Type species. 
of the U.S.A. 



Leptaena subquadrata Hall, 1 883, from the Caradoc 



PLATE 2 

Figs 1-4 Furcitellinae gen. et sp. indet. 1, 2, 4. Sample 100. Anderkenyn-Akchoku section; 1, 2, BC 57382, dorsal radial ornament and exterior, x 5. x 2; 
4, BC 57384, ventral internal mould, x 2. 3, Sample 1041a, Burultas Valley, BC 57383, ventral exterior, x 2.5. 

Figs 5-9 Glyptomena onerosa Popov. 5, Sample 7613, Akchoku Mountain, Kujandysai section, BC 57388, dorsal valve ornament, x 3. 6. Sample 8229, 
Buldukbai-Akchoku section, west side of Kopalysai, BC 57385. ventral internal mould, x 2. 7, Sample 8235, Anderkenyn-Akchoku section, BC 57386, 
ventral internal mould, x 3. 8, Sample 100b, Anderkenyn-Akchoku section, CNIGR Museum, dorsal internal mould, x 4. 9. Sample 843, Anderkenyn- 
Akchoku section, BC 57387, dorsal exterior, x 1.5. 

Figs 10-20 Teretelasmella chugaevae sp. nov. Anderkenyn-Akchoku section. 10-13, 19, 20, Sample 100; 10-13, 19, BC 57391, conjoined valves, anterior, 
posterior, dorsal, lateral and ventral views, x 1.5; 20. BC 57390, conjoined valves, posterior view, x 2. 14-18, Sample 626. BC 57392, conjoined valves, 
holotype, dorsal, posterior, lateral, ventral and anterior views, x 1.5. 



36 



L.E. POPOV, L.R.M. COCKS AND I.F. NIKITIN 




UPPER ORDOVICIAN BRACHIOPODS FROM KAZAKHSTAN 



37 



Cristiania egregia Popov, 1985 PI. 3, figs 2-6, 8 

1985 Christiania egregia Popov: 60, pi. 2, figs 7-1 1 . 

HOLOTYPE. CNIGR 25/11989, dorsal internal mould from the 
Anderken Formation, Sample 1018, 7 km south-west of Karpkuduk 
Well, Kotnak Mountains. 

Material. Eight pairs of conjoined valves. 17 ventral and 30 
dorsal valves from Samples 100 (BC 56875-81), 843, 626 (BC 
56869-70), 8214 (BC 56871-2), Anderkenyn-Akchoku section; 
Samples 7613, 628 (BC 56587, 56886-89), Kujandysai section; 
Sample 1041a (BC 57396-98), Burultas Valley; Sample 1024b, east 
side of Karatal River near Sorbulak well; Sample 1018 (BC 56873- 
74), 7 km south-west of Karpkuduk well, Kotnak Mountains. 

Description. Shell concavoconvex, strongly elongated, suboval 
in outline, about 160% as long as wide with maximum width about 
two-thirds valve length from the hinge line. Cardinal extremities 
subrectangular, usually slightly alate. Anterior commissure broadly 
uniplicate. Ventral valve moderately convex in lateral profile with 
maximum thickness at about one-third valve length, about 20% as 
thick as long. Ventral interarea strongly apsacline to orthocline with 
a large convex pseudodeltidium perforated apically by a minute, 
circular foramen. Very weak ventral sulcus originating at the umbonal 
area. Dorsal valve gently concave with hypercline interarea and 
complete, convex chilidium. Radial ornament finely parvicostellate, 
rarely preserved. 

Ventral interior with strong, transverse teeth and low, widely 
diverging dental plates. Muscle field weakly impressed, bilobate, 
with short, linear adductor scars bisected by a fine median ridge. 
Dorsal interior with bilobed cardinal process, low, curved socket 
ridges subparallel to the hinge line and two pairs of strong side septa. 
Dorsal median septum very fine, about 61% as long as the valve. 
Adductor scars bordered anteriorly by strong muscle bounding 
ridges curved posteriorly. 

Discussion. Detailed discussion of this species was provided by 
Popov (1980: 61). Among the Kazakh species it is most similar to 
Christiania tortuosa Popov, 1980 from the Lidievka Formation 
(Llandeilo-LowerCaradoc) of north-central Kazakhstan, but differs 
in having a weak ventral sulcus, very fine radial ornament and a long 



dorsal median ridge extending far beyond the mid-valve. There are a 
large number of nominal Christiania species known globally, and 
the genus as a whole is due for revision. 

Christiania aff. sulcata Williams, 1962 PI. 3, figs 7, 9-17 

Material. Three pairs of conjoined valves, 23 ventral and 10 
dorsal valves from Samples 8223 (BC 56585-6), 8223a, 8223b (BC 
57399, 57401), Anderkenyn-Akchoku section; Sample 8215 (BC 
56583^1), west side of Ashchisu River; Samples 628 (=K- 1 07/1 970) 
(BC 56593), 2538 (BC 56579, 81, 89), 8217 (BC 57400), 8220 (BC 
56592), Kujandysai section. 

Description. Shell concavoconvex, elongate and subtrapezoidal 
in outline, as long as wide with maximum width at three-quarters 
valve length. Hinge line about 85% of maximum shell width. Cardi- 
nal extremities near right-angled and slightly alate. Lateral 
commissures near straight, slightly diverging anteriorly. Anterior 
commissure gently uniplicate. Ventral valve strongly convex in 
profile with maximum thickness at about one-third valve length. 
Ventral interarea strongly apsacline to near orthocline with narrow 
convex pseudodeltidium perforated apically by a minute rounded 
foramen. Lateral sides of the valve steep inclined near right angle 
towards the commissural plane. A shallow sulcus v-shaped in cross- 
section originating near the umbo and flanked by two distinct 
plications rounded in cross-section. Dorsal valve moderately con- 
cave with low and narrow median fold. Dorsal interarea hypercline 
with a convex chilidium. Shell surface finely and equally 
parvicostellate with 1 6 to 1 8 parvicostellae along the anterior margin 
of mature specimens. 

Ventral interior with small, bilobed muscle field, fine teeth and 
rudimentary dental plates. Dorsal interior with a double cardinal 
process, low, curved socket ridges, thin median septum extending 
somewhat anterior of mid- valve and two pairs of strong side septa. 

Measurements. (457/12375) ventral valve, L=8.2, Iw=5.7, 
W=6.8, T=3.8, Sw=3.2; 458/12375), dorsal valve, L=5.6, Iw=5.2, 
W=5.3, Sw=2.2 

DISCUSSION. These specimens resemble Christiania sulcata 
Williams, 1962, from the Stinchar Limestone (Upper Llandeilo- 



PLATE 3 

Fig. 1 Limbimurina sp. Sample 626, Anderkenyn-Akchoku section, BC 57395. ventral exterior, x 1.5. 

Figs 2-6, 8 Christiania egregia Popov. 2, 3, Sample 1041a, Burultas Valley, BC 57396, lateral and ventral views of exterior, x 2. 4, Sample 1018, area 7 
km west of Karpkuduk well, Kotnak Mountains, south Betpak-Dala, CNIGR 22/1 1989, latex cast of dorsal interior, x 2. 5, Sample 1041a, BC 57397, 
dorsal valve interior, x 2. 6, Sample 1018, CNIGR 23/1 1989, ventral internal mould, x 2. 8, Sample 1041a, BC 57398, conjoined valves dorsal view 
showing interareas, x 4. 

Figs 7, 9-17 Christiania aff. sulcata Williams. 7, Sample 628, west side of Kujandysai, BC 56593, dorsal exterior, x 4. 9, Sample 8223b, Anderkenyn- 
Akchoku section, BC 57399, ventral exterior, x 3. 10, Sample 8217, BC 57400, ventral exterior, x 3. 11-16, Sample 2538, Akchoku Mountain, 
Kujandysai section; 11, BC 56589, dorsal external mould, x 3; 12-14, BC 56579, conjoined valves, posterior view, x 8, dorsal and ventral views, x 6; 15, 
16, BC 56581, dorsal internal mould and latex cast, x 5. 17, Sample 8223b, BC 57401, ventral internal mould, x 3. 

Figs 18-23, 25 Foliomena prisca sp. nov. 18, 19, 21-23, Sample 8255, Anderkenyn-Akchoku section; 18, BC 57402, latex cast of dorsal exterior, x 4; 19, 
BC 57403, latex cast of dorsal exterior, x 4; 21, BC 57404. latex cast of dorsal exterior, x 4; 22, 23, BC 57405, holotype, latex cast and dorsal internal 
mould, x 4. 20, Sample 100, Anderkenyn-Akchoku section, BC 57407, dorsal exterior, x 4. 25, Sample 8217, Kujandysai section, BC 57408, ventral 
internal mould, x 4. 

Fig. 24 Kassinella (Kassinella)? sp. Sample 2531, Anderkenyn-Akchoku section, BC 56497, dorsal internal mould, x 8. 

Figs 26-34 Craspedelia tata Popov. 26-28, Sample 626, Anderkenyn-Akchoku section, BC 57409, conjoined valves, dorsal, lateral and ventral views, x 4. 
29-30, Sample 100, BC57410, dorsal exterior and anterior views, x 3. 31, Sample 8238, CNIGR 9/12361, ventral exterior, x 2. 32-34, Sample 626; 32, 
34, BC 5741 1, dorsal anterior and exterior view, x 3; 33, BC 57412, conjoined valves, dorsal view, x 5. 

Fig. 35 Isophragma imperator Popov, Sample 1018, 7 km southwest of Karpkuduk well, Kotnak Mountains, south Betpak-Dala. CNIGR 22/1 1522, latex 
cast of dorsal interior, x 2. 

Figs 36-40 Acculina kulanketpesica sp. nov. 36, Sample 8231^10. Buldukbai-Akchoku section, BC 57416, ventral exterior, x 2. 37^M), Sample 1041a, 
Burultas Valley; 37, BC 57415, posterior view of ventral and dorsal interareas, x 5; 38, BC 12903, ventral interior, x 2; 39, 40, BC 12904, holotype, 
conjoined valves, ventral and dorsal views, x 3. 



38 



L.E. POPOV, L.R.M. COCKS AND I.F. NIKITIN 



Table 9 Measurements of ventral valves of Christiania aff. sulcata 
Williams. Parastrophina-Kellerella Association, Anderkenyn-Akchoku 
and Kujandysai sections. 





Lv 


W 


Iw 


Sw 


Lv/W 


Iw/W 


Sw/W 


N 


9 


9 


6 


5 


9 


6 


5 


X 


6.5 


5.6 


4.8 


2.5 


115.0% 


87.8% 


42.9% 


s 


1.91 


1.15 


0.85 


0.73 


16.1 


6.4 


12.4 


MIN 


4.0 


4.4 


3.9 


1.7 


90.9% 


82.8% 


23.0% 


MAX 


9.4 


7.4 


6.3 


3.2 


147.5% 


100.0% 


54.2% 



Lower Caradoc) of Girvan, Scotland, but differ in having an elongate 
subtrapezoidal shell outline and two rounded plications flanking the 
ventral sulcus. A characteristic feature of the Kazakh specimens is 
the absence of high ridges bordering the dorsal adductor field 
anteriorly, which also appear to be absent in C. sulcata (Williams 
1962: pi. 18, fig. 36). Christiania aff. sulcata also differs from C. 
egregia in having a stronger ventral median sulcus which is v-shaped 
in cross-section, a distinctive dorsal median fold and significantly 
smaller size. 



Family FOLIOMENIDAE Williams, 1965 
Genus FOLIOMENA Havh'cek, 1952 

Type species. Strophomena folium Barrande, 1879, from the 
Kralfiv Dvur Formation ( Ashgill) of Bohemia. 

Foliomena prisca sp. nov. PI. 3, figs 18-23, 25 

Etymology. After priscus, Latin - old. 

Holotype. BC 57405, PI. 3, figs 22, 23, a dorsal valve, from the 
Anderken Formation, Sample 8255, Anderkenyn-Akchoku section. 

Material. 2 pairs of conjoined valves, 4 ventral and 3 dorsal 
valves from Samples 100 (BC 57407), 2531, 8221 (BC 56890), 
8223b, 8251, 8255 (BC 57402-06, 08), Anderkenyn-Akchoku sec- 
tion; 628, 2538, 8217, Kujandysai section. 

Description. Shell flat and gently resupinate, transverse, 
subrectangular in outline, about 60% as long as wide, with maximum 
width at hinge line. Cardinal extremities near right angled. Anterior 
commissure rectimarginate. Ventral valve gently convex in the pos- 
terior half and weakly concave anteriorly. Ventral interarea apsacline 
with a minute, apical pseudodeltidium. Dorsal valve with lateral 
profile flat to slightly concave in the posterior half and convex 
posteriorly, with maximum thickness at about three-quarters valve 
length in mature specimens. Dorsal interarea linear, anacline. with 
separate chilidial plates flanking narrow notothyrium. Radial orna- 
ment of fine capillae rarely preserved. Concentric ornament of 
numerous slightly uneven fine rugellae. 

Ventral interior with delicate teeth lacking dental plates and an 
open, weakly impressed muscle field. Dorsal interior with a small 
bilobed cardinal process in the low notothyrial platform and thin 
widely diverging socket ridges. Thin median septum, about half the 
length of the pair of strong curved side septa. 

Measurements. (463/12375) ventral valve, L=6.0, W=l 1.7; (464/ 
12375) ventral valve, L=6.4, W=11.4; (466/12375) dorsal valve, 
L=4.3,W=7.2; (468/1 2375) dorsal valve, L=4.6,W=8.5; (469/1 2375) 
dorsal valve, L=3.6, W=6.3. 

Discussion. Cocks & Rong ( 1988:65) discussed the variation in 
the ornament of Foliomena and noted that, although all the shells 
from the type locality in Bohemia were devoid of radial ornament. 



occasional costae or costellae can be present sporadically in some 
populations. The presence of fine capillae is a key feature in our new 
species, as is the resupinate shell shape and the separate chilidial 
plates, although the features within the interarea are poorly known in 
Foliomena folium. It is difficult to make a precise comparison of our 
shells with F. inelegans Fu, 1982 from the Pingliang Formation 
( Upper Caradoc) of North China, because of inadequate information 
on the interior of the latter species. From F.jielingensis, described by 
Zeng (1987) from the Miapo Formation (Lower Caradoc) of the 
Yangtze Gorge area, south China, F. prisca differs in the presence of 
the fine ornament, the resupinate shell shape and in the absence of the 
ventral internal tuberculae seen in F.jielingensis. 



Superfamily PLECTAMBONITOIDEA Jones, 1928 

Family PLECTAMBONITIDAE Jones, 1928 

Subfamily TAPHRODONTINAE Cooper, 1956 

Genus ISOPHRAGMA Cooper, 1956 

TYPE SPECIES. Isophragma ricevillense Cooper, 1956. from the 
Lower Caradoc of Tennessee, U.S.A. 

Isophragma imperator Popov, 1980 PI. 3, fig. 35 

1980 Isophragma imperator Popov: 147, pi. 2, figs 8-12. 

Holotype. CNIGR 25/1 1523, from Sample 1018. 7 km south-west 
of Karpkuduk Well, Kotnak Mountains. 

Material. One pair of conjoined valves, 36 ventral and 34 dorsal 
valves from Sample 1018. 

Discussion. Popov (1980) provided detailed description and dis- 
cussion of this species. 



Family BIMURIIDAE Cooper, 1956 
Genus CRASPEDELIA Cooper, 1956 

TYPE SPECIES. Craspedelia marginata Cooper, 1956: 773, pi. 2 13, 
figs. 1-20, from the Pratt Ferry Formation (Landeilo), Alabama, 
U.S.A. 



Craspedelia lata Popov, 1980 



PL 3, figs 26-34 



1 980 Craspedelia lata Popov: 55, pi. 1 7, figs 6-9. 

1986 Craspedelia tata Popov; Kolobova & Popov: pi. 1 , fig. 9. 

Holotype. CNIGR 8/ 1 1 098, from the Lidievka Formation (Lower 
Caradoc), Belyi Kardon, north-central Kazakhstan. 

Material. 25 pairs of conjoined valves, 18 ventral and 5 dorsal 
valves from Samples 100 (BC 56900-03, 57410, 57597) (=K98/ 
1970), 626 (BC 56532, 56907-10, 57409, 57411-12), 8223a (BC 
56935), 8223b, Anderkenyn-Akchoku section; Samples 8214 (BC 
56925-30, 56932-33, 57598), 8215b (BC 56931), west side of 
Ashchisu River; Samples 628 (BC 56911), 2538 (BC 56917-24), 
Kujandysai section; Sample 8231^40, Buldukbai-Akchoku; Sample 
948 (BC 56912-16), Tesik River. 

Description. Shell smooth, concavoconvex posteriorly, strongly 
geniculated ventrally with a trail up to 8 mm long which curves back 
postero-ventrally; semielliptical in outline, about 80% as long as 
wide, with maximum width slightly anterior to hinge line. Cardinal 
extremities rounded. Anterior commissure uniplicate. Ventral valve 
strongly convex posteriorly with maximum thickness about one- 
third anteriorly. Narrow and shallow sulcus anterior to the 



UPPER ORDOVICIAN BRACHIOPODS FROM KAZAKHSTAN 



39 



geniculation. Dorsal valve strongly concave with low, planar interarea 
and notothyrium covered by joined apical chilidial plates. Weak 
median fold originating anterior to the geniculation. Shell surface 
smooth with fine growth lines. Ventral interior with strong teeth and 
vascula media subparallel. Dorsal interior with simple, undercut 
cardinal process, divided bema, low median ridge and a pair of 
slightly diverging side ridges bisecting the bema. 

DISCUSSION. This species differs from Craspedelia marginata 
Cooper ( 1 956) and C. gabata Williams ( 1 962: 1 79), from the Lower 
Ardwell Formation (Middle Caradoc) of Girvan, in its high (up to 8 
mm) ventrally directed trail with a well-defined ventral sulcus and 
dorsal median fold. It differs from C. intonsa Potter, 1991, from 
Member 1 of the Gregg Ranch Unit (Llandeilo), of California. USA, 
in having a larger shell, up to 12 mm long, with a less transverse 
outline and a high trail, which exceeds the thickness of the ventral 
valve in mature specimens. 



Family LEPTELLINIDAE Ulrich & Cooper 1936 

Subfamily LEPTELLININAE Ulrich & Cooper 1936 

Genus ACCULINA Misius, in Misius & Ushatinskaya,1977 

TYPE SPECIES. Acculina acculica Misius in Misius & Ushatinskaya, 
1977, from the Tabylgaty Formation (Lower Caradoc: gracilis Zone), 
Moldo-Too Range, North Kirgyzstan. 

Acculina kulanketpesica sp. nov. PI. 3, figs 36-40, PI. 4, 
figs 1-5 

Etymology. After Kulanketpes ('donkey cannot escape' in 
Kazakh) Valley on the way from Lake Balkhash to the type locality. 

Holotype. BC 12904, PI. 3, figs 39, 40, conjoined valves, from 
the Anderken Formation, Sample 1041a, Burultas section. 

MATERIAL. 50 pairs of conjoined valves, 13 ventral and 10 dorsal 
valves from Samples 100 (=K98/1970) (BC 56493-5, 56485-90, 
BC 57414), 620 (BC 56937-39), 626, 8128, Anderkenyn-Akchoku 
section; Sample 823 1-^0 (BC 57416, 18), Buldukbai-Akchoku; 
Sample 1041a (BC 12900-7,57413, 15, 19), Burultas Valley; Sam- 
ples 85258 (BC 56491-2, 56941-6), 2538 (BC 56501-02), 
Kujandysai. 

Description. Shell resupinate, transverse, semielliptical in out- 
line, about 70% as long as wide and 35% as thick as long. Cardinal 
extremities near right-angled or slightly acute. Anterior commissure 
rectimarginate. Ventral valve gently to moderately concave in the 
anterior half and slightly convex posterior to the mid- valve. Ventral 
interarea apsacline with a narrow, convex pseudodeltidium. Dorsal 
valve flat and gently sulcate between the umbo and mid-length, 
becoming moderately convex anteriorly. Interarea anacline with a 
narrow, convex chilidium completely covering the notothyrium. 
Radial ornament unequally parvicostellate with 7 accentuated costae 
in the umbonal area and 27-30 accentuated costellae along the 

Table 10 Basic statistics of complete shells of Acculina kulanketpesica 
sp. nov. from Sample F- 104 la. Burultas valley. 





Lv 


Ld 


W 


T 


Lv/W 


LdAV 


T/Lv 


N 


6 


5 


6 


5 


6 


5 


5 


X 


15.5 


14.7 


22.6 


5.4 


68.9% 


65.6% 


34.5% 


S 


0.89 


0.92 


0.85 


1.13 


1.9 


2.1 


7.0 


MIN 


14.0 


13.5 


21.3 


3.8 


65.7 


63.4 


24.1 


MAX 


16.4 


16.1 


23.5 


7.0 


71.2% 


68.5% 


42.9% 



anterior and lateral margins in full grown specimens. Interspaces 
between the accentuated costellae occupied by fine parvicostellae 
about 7-10 per mm along the anterior margin. Fine, closely spaced 
growth lamellae form comae crossed by accentuated ribs in the 
anterior half of the shell. 

Ventral valve with strong teeth with central grooves; lacking 
dental plates. Ventral muscle field small, rounded pentagonal with 
strongly impressed diductor scars completely divided by slightly 
shorter, narrow, triangular adductor scars. Ventral mantle canals 
saccate with slightly diverging vascula media. Dorsal valve interior 
with trifid cardinal process situated on the low notothyrial platform. 
Socket ridges low, widely diverging and slightly incurved posteriorly. 
Dorsal adductor muscle field subquadrate, bordered laterally by low, 
subparallel ridges. Median ridge high and strongly thickened, ex- 
tended anteriorly to the mid-valve, merging with high diaphragm 
bordering the lophophore platform. Dorsal mantle canals lemniscate. 

Discussion. This species also occurs in Betpak Dala, Kazakhstan 
(the locality in Nikitin & Popov 1996). It differs from Acculina 
acculica Misius {in Misius & Ushatinskaya 1977: 1 14) in its signifi- 
cantly larger shell, strongly convex anteriorly transverse profile of 
the dorsal valve and numerous comae (PL 3, fig. 39) in the anterior 
part of full grown specimens. It is also similar to A. villosa Nikitina 
( 1 985 : 24) from the Rgaity Formation ( Llandeilo-Lower Caradoc) of 
the Kendyktas Range, south Kazakhstan, in the development of 
comae, but can be distinguished in having a well-defined peripheral 
rim in the ventral valve, a somewhat smaller dorsal lophophore 
platform, which has a subrectangular, not a flabellate outline, and a 
pair of fine transmuscle ridges dividing the anterior and posterior 
adductors. Acculina is the only plectambonitoid in the Anderken 
Formation to possess comae, and its exterior is also normally covered 
by encrusting girvanellid algae. Two specimens (BC 56502-3, PI. 4, 
figs 7, 8) from the Kujandysai section from Sample 2538 differ from 
other Acculina in having a geniculate shell with the lophophore 
platform less than half the valve length, and probably represent a 
separate species. 



Genus DULANKARELLA Rukavishnikova, 1956 

Type SPECIES. Dulankarella magna Rukavishnikova 1956, from 
the Dulankara Formation (late Caradoc), Kazakhstan, Chu-Ili Range. 
Kazakhstan. 

Dulankarella larga sp. nov. PI. 4, figs 9-25, PI. 5, figs 1-3 

ETYMOLOGY. After largus, Latin - rich. 

Holotype. BC 57421, PI. 4, figs 9, 10, a dorsal valve interior from 
Anderken Formation, Sample 8231-40, Buldukbai-Akchoku sec- 
tion. 

Material. 86 pairs of conjoined valves, 3 ventral and 2 dorsal 
valves, from Samples 100 (=K98/1970) (BC 56514, 16-18, 25-29, 
57422, 24, 26), 626 (BC 56522-4), 8223a (BC 56511, 56975), 
Anderkenyn-Akchoku; Sample 8231^10 (BC 57421, 23), Buldukbai- 
Akchoku; Sample 8228 (BC 56986), east side of Kopalysai River; 
Samples 1041a (BC 56512, 19-21, 56947-57), 818a, 1041a (BC 
56967, 57425), Burultas Valley. 

DESCRIPTION. Shell strongly concavoconvex, transverse and 
semielliptical in outline, on average 73% as wide as long, with 
maximum width at the hinge line and 45% as thick as long. Cardinal 
extremities slightly acute. Anterior commissure weakly sulcate. 
Ventral valve strongly and evenly convex in lateral profile, subcarinate 
posteriorly in transverse profile. Ventral interarea low, planar, anacline 



40 



L.E. POPOV, L.R.M. COCKS AND I.F. NIKITIN 




UPPER ORDOVICIAN BRACHIOPODS FROM KAZAKHSTAN 



41 



Table 11 Basic statistics of complete shells of Dulankarella larga sp. 
nov. from Sample F- 104 la, Burultas valley. 





Lv 


W 


T 


Lv/W 


T/Lv 


N 


9 


9 


7 


9 


7 


X 


16.9 


24.6 


7.7 


69.7% 


46.1% 


s 


2.10 


3.57 


1.19 


10.2 


3.2 


MEN 


13.5 


19.1 


5.7 


51.9% 


41.5% 


MAX 


18.8 


31.6 


9.2 


81.3% 


50.0% 



with delthyrium partly covered by a convex pseudodeltidium. Dorsal 
valve strongly concave and slightly geniculate anteriorly. Dorsal 
interarea hypercline with chilidial plates joined apically. Sulcus 
broad and shallow, originating near mid-valve. Radial ornament 
unequally parvicostellate with 5-7 accentuated ribs originating at 
the umbo and two or three generations of accentuated costellae, 
totaling 31-38 in number in full grown specimens. Parvicostellae 
between accentuated ribs very fine and closely spaced, about 12-16 
per mm along the anterior margin. 

Ventral valve interior with strong teeth supported by short den- 
tal plates, thickened at the base. Muscle field bilobed with strongly 
impressed, rounded subrhomboidal diductor scars separated by 
short, elongate subtriangular adductor scars. Paired nodose swell- 
ings anterolateral to the muscle field. Ventral mantle canals saccate 
with short, divergent vascula media. Dorsal interior with trifid 
cardinal process bearing a strong, ridge-like median lobe and up to 
6 fine ridges on the lateral lobes. Socket ridges narrow, widely 
diverging. Median septum strongly raised and thickened anteriorly 
with the maximum height at the point of junction with the outer 
boundary of the lophopore platform accentuated by geniculation 
of the valve. 

Discussion. This species resembles the later Dulankarella magna 
Rukavishnikova ( 1956: 139) in size and transverse profile, but differs 
in having a finer radial ornament with 5-7 strongly accentuated 
primary ribs and a characteristic rounded subrhomboidal outline of 
the ventral diductor scars, which only slightly touch each other 
anteriorly to the adductor scars. The differences from Dulankarella? 
partita Percival (1979b: 103) are in having a less transverse outline, 
evenly convex ventral valve, the ventral muscle field lacking a 
median ridge, and the rhomboidal outline of the ventral diductor 
scars. 



Kajnaria rugosa sp. nov. PL 5, figs 6-18 

Etymology. After rugosus, Latin - wrinkled. 

HOLOTYPE. BC 5655 1 , PI. 5, figs 12, 1 3, a dorsal internal mould 
from Sample 628, Kujandysai section. 

Material. 9 pairs of conjoined valves and three ventral valves 
from Samples 100 (BC 56545-47, 57436, 37), 626 (BC 56548), 
8223a, Anderkenyn-Akchoku section; Sample 628 (=K107/1970) 
(BC 56551, 57439), Kujandysai section; Sample 1041a (BC 56543, 
44), Burultas Valley: Sample 816 (BC 56549-50, 57438), Alakul 
lake. 

Description. Shell strongly concavoconvex, semielliptical in out- 
line, about 67-82% as long as wide with maximum width at the 
hinge line. Cardinal extremities acute and slightly alate in adult 
specimens. Ventral valve strongly convex in lateral profile, weakly 
geniculate in some specimens, with maximum thickness posterior to 
mid-length. Relatively weak umbo. Interarea anacline with narrow 
convex pseudodeltidium. Dorsal valve moderately and evenly con- 
vex with hypercline interarea. Notothyrium completely covered by 
convex chilidium. Radial ornament unequally parvicostellate with 
5-7 primary accentuated ribs and two to three generations of accen- 
tuated costellae, with the interspaces between them covered by fine 
closely-spaced parvicostellae, about 7-14 per mm. Concentric orna- 
ment of up to 9 undulated rugellae in the posterior half of both valves. 
Ventral interior with strong teeth lacking dental plates. Ventral 
muscle field small, strongly supported by raised muscle bounding 
ridges which merge centrally and anteriorly. A further pair of strong 
curved ridges originate posteriorly at one-third of the length of the 
hinge line on each side of the valve, and curve anterolaterally to form 
a w-shaped structure which terminates near the end of the bounding 
ridges at about quarter valve length. Mantle canals saccate with 
vascula media subparallel in the proximal part and diverging 
anteriorly. Dorsal interior with erect trifid cardinal process fused 
anteriorly to a strong median ridge. Sockets large. The median ridge 
merges near the mid-length with a high subperipheral rim bordering 
the lophophore platform. 

DISCUSSION. This species differs from Kajnaria derupta in the 
twice as large shell size and the well defined concentric rugellae in 
the posterior half of the valves. 



Genus KAJNARIA Nikitin & Popov, 1984 

TYPE SPECIES. Kajnaria derupta Nikitin & Popov in Klenina et al. 
(1984), from the Bestamak Formation (Lower Caradoc), Chingiz 
Range, Kazakhstan. 



Genus MABELLA Klenina, 1984 

TYPE SPECIES. Leptellina (Mabella) semiovalis Klenina in Klenina 
et al. (1984), from the Taldyboy Formation, Dulankara Regional 
Stage (Upper Caradoc), Chingiz Range, Kazakhstan. 



PLATE 4 

Figs 1-5 Acculina kulanketpesica sp. nov. 1, Sample 1041a, Burultas Valley, BC 57413, dorsal interior, x 2. 2, Sample 8137, Anderkenyn-Akchoku 
section, BC 57417, latex cast of dorsal interior, x 3. 3, Sample 100, Anderkenyn-Akchoku section, BC 57414, dorsal exterior view of conjoined valves, x 
2. 4, Sample 1018, area 7 km S W of Karpkuduk well, Kotnak Mountains, ventral internal mould, x 3. 5, Sample 85258, east of Uzunbulak River, BC 
56491, ventral internal mould, x 2. 

Fig. 6 Glyptambonites sp., Sample 628 (=K-107/70), west side of Kujandysai, BC 56510, ventral exterior, x 3. 

Figs 7, 8 Acculina sp. Sample 2538, Akchoku Mountain, Kujandysai section. 7, BC 56502, latex cast of dorsal interior, x 2. 8, BC 56501. ventral internal 
mould, x 1.5. 

Figs 9-25 Dulankarella larga sp. nov. 9, 10, 16, Sample 823 1^10 , Buldukbai- Akchoku section, west side of Kopalysai; 9, 10, BC 5742 1 , holotype. 
dorsal interior, x 2.7; cardinal process and socket plates, x 5; 16, BC 57423, ventral internal mould, x 2. 11-13, 19-21, 24, 25, Sample 1041a, Burultas 
Valley; 11-13, conjoined valves, lateral, ventral and dorsal views, x 2; 19-21. BC 57425, conjoined valves, ventral, dorsal and lateral views, x 2; 24, 
Sample 1041a, BC 56512, conjoined valves, posterior view showing interareas. x 6: 25, BC 56967. dorsal view of conjoined valves, x 2. 14, 15, 17, 18, 
22, 23, Sample 100, Anderkenyn-Akchoku section; 14, BC 56514, incomplete dorsal valve interior, x 3: 15, BC 57422, ventral internal mould, x 2; 17, 
18, BC 57424, conjoined valves, ventral and dorsal views, x 1.5; 22, 23, BC 57426, ventral valve exterior, anterior and ventral views, x 2. 



42 



L.E. POPOV, L.R.M. COCKS AND I.F. NIKITIN 




UPPER ORDOVICIAN BRACHIOPODS FROM KAZAKHSTAN 



43 



Species included. Leptellina (Mabella) semiovalis Klenina, in 
Klenina e? a/. 1984: 69, pi. 5, figs. 1,3, 4; pi. 9, figs 4, 7 (^Leptellina 
(Mabella) obtusa Klenina, in Klenina etal. 1984: 71, pi. 5, figs 5, 6; 
pi. 6, fig. 2; ^Leptellina (Mabella) incurvata Klenina, in Klenina et 
al. 1984: 72, pi. 5,fig. 2), Upper Caradoc, beds tb n v of Taldyboi 
Formation, Chingiz Range, Kazakhstan; Leptellina ? conferta Popov, 
1985: 56, pi. 2, figs 1-6, Lower Caradoc, Anderken Formation, Chu- 
Ili Range; Leptelloidea multicostata Rukavishnikova, 1956: 132; 
Wiradjuriella halis Percival, 1991: 138, fig. 1 2A-Z, Aa-Al, Upper 
Caradoc, New South Wales, Australia; Leptellina sp., Percival, 1 979b, 
Ordovician, Goonumbla Volcanics, New South Wales, Australia. 

DISCUSSION. Mabella differs from Leptellina in the distinctive 
median septum which enlarges anteriorly, and is sometimes bifurcat- 
ing and tubular. However, Wiradjuriella, from the Caradoc of Australia 
(Percival 1991), has this same structure as Mabella and can be 
considered congeneric with it (Cocks & Rong 2000). 

Mabella conferta (Popov, 1985) PI. 5, figs 19-29 

1985 Leptellina! conferta Popov: 56, pi. 2, figs 1-6, text-figs 1 , 2. 
1991 Wiradjuriella conferta (Popov) Percival: 140. 

Holotype. CNIGR 17/11989, ventral internal mould from the 
Anderken Formation, Sample 100b, Anderkenyn-Akchoku section. 

Material. 35 pairs of conjoined valves, 183 ventral and 66 dorsal 
valves from Samples 100, 100b, 843, 8128a, 8128b, 8137 (BC 
56982-84), Anderkenyn-Akchoku section; Sample 761 3, Kujandysai 
section; Samples 110, 8229, 8230 (BC 57440), 8257, Buldukbai- 
Akchoku; Sample 8228 (BC 57441, 43^16), east side of Kopalysai 
River; Samples 818a (BC 57442), 1041a (BC 56978-81), Burultas 
Valley; Samples 1018, 1018a, 7 km south-west of Karpkuduk well, 
Kotnak Mountains. 

DESCRIPTION. Shell concavoconvex, transverse, semioval in out- 
line, length about three-quarters of the width, with maximum width 
at the hinge line. Anterior commissure rectimarginate. Ventral valve 
strongly convex in transverse and lateral profiles with the maximum 
thickness slightly posterior to mid-length. Planar strongly apsacline 
interarea and small, convex pseudodeltidium. Dorsal valve gently 
concave to almost flat, with a planar, anacline interarea and disjunct 
chilidial plates. Radial ornament very fine, unequally parvicostellate, 
with up to five accentuated parvicostellae per 3 mm along the 
anterior margin of mature specimens. 

Ventral interior with strong teeth lacking dental plates. Cordate 
muscle field with short, ridge-like adductor scars completely sepa- 
rating strongly impressed diductor scars. Strong, slightly divergent, 
saccate mantle canals. Dorsal interior with low, trifid cardinal process 
facing posteriorly, short socket ridges subparallel to the hinge line. 



Lophophore platform about 90% valve length and 87% as wide as 
maximum valve width, bordered by a high, ridge-like rim divided 
medially. High median septum about three-quarters as long as the 
valve, not joined anteriorly with the subperipheral rim. 

DISCUSSION. This species was originally assigned to Leptellina 
and later referred by Percival ( 1991 ) to Wiradjuriella. Percival also 
listed and discussed the differences between the various species of 
the genus, which is so far known only from Australia and Kazakhstan. 

Genus SHLYGINIA Nikitin & Popov, 1983 

TYPE SPECIES. Shlyginia declivis Nikitin & Popov, 1983, from the 
Andriushino Formation, Tselnograd Regional Stage (Llandeilo- 
Lower Caradoc), north-central Kazakhstan. 

Discussion. The affinities of Shlyginia and its differences from 
Dulankarella were discussed by Nikitin & Popov (1996). 

Table 12 Measurements of complete shells of Shlyginia fragilis 
(Rukavishnikova) Sample 8228 and 8257 from Kopalysai section. 



Lv 



W 



LAV 



T/L 



N 


16 


16 


16 


16 


16 


X 


10.4 


16.3 


4.2 


64.0% 


40.4% 


s 


1.04 


1.34 


0.54 


6.2 


4.5 


MIN 


8.5 


13.6 


2.9 


50.0% 


31.2% 


MAX 


12.2 


19 


5.1 


72.1% 


51.8% 



Table 13 Measurements of ventral valves of Shlyginia fragilis 
(Rukavishnikova) Sample 8228 from Kopalysai section. 



Lv 



W 



Ml 



Mw 



LvAV 



M//L Ml/Mw 



N 


6 


6 


6 


6 


6 


6 


6 


X 


9.4 


12.8 


3.6 


4.3 


75.2% 


37.6% 


83.5% 


s 


2.47 


3.61 


1.13 


1.05 


11.2 


4.5 


16.3 


MIN 


5.3 


5.8 


1.8 


2.6 


60.0% 


30.5% 


67.9% 


MAX 


12.8 


15.5 


5.3 


5.6 


91.4% 


41.4% 


110.4% 



Table 14 Measurements of dorsal valves of Shlyginia fragilis 

(Rukavishnikova) Sample 8228 from Kopalysai section and sample 8229 
from Buldukbai-Akchoku. 





Ld 


W 


SI 


BB1 


BBw 


Ld/W 


Sl/L 


BBwAV 


N 


6 


6 


3 


6 


6 


6 


3 


6 


X 


8.1 


12.1 


5.5 


1.2 


3.8 


67.7% 


79.2% 


31.7% 


s 


1.87 


2.55 


1.25 


0.42 


0.93 


13.2 


8.6 


8.5 


MIN 


6.2 


7.8 


4.3 


0.6 


2.8 


49.7% 


69.4% 


23.6% 


MAX 


10.8 


14.8 


6.8 


1.8 


5.0 


82.1% 


84.4% 


43.6% 



PLATE 5 

Figs 1-3 Dulankarella larga sp. nov. 1, 2, Sample 100, Anderkenyn-Akchoku, BC 57427, ventral exterior and lateral views, x 2. 3, Sample 1041a, 
Burultas Valley, BC 56513, interareas of conjoined valves, x 8. 

Figs 4, 5 Chonetoidea sp. Sample 8255, Anderkenyn-Akchoku section, BC 56537, external and internal moulds of conjoined valves, x 8. 

Figs 6-18 Kajnaria rugosa sp. nov. 6, unnamed Lower Caradoc formation, about 4 km south-west of Lake Alakul, Sample 816, BC 57438, ventral internal 
mould, x 1.5. 7-11, 14-16, Sample 100. Anderkenyn-Akchoku; 7-10, BC 57436, conjoined valves, lateral, posterior, ventral and dorsal views, x 2; 11, 14- 
16, BC 57437, conjoined valves, lateral, posterior, ventral and dorsal views, x 2. 12, 13, 17, 18, Sample 628, west side of Kujandysai River; 12, 13, BC 
5655 1 , latex cast and dorsal internal mould, holotype, x 2; 17, 18, BC 57439, ventral internal mould, oblique posterior and oblique anterior views, x 2. 

Figs 19-29 Mabella conferta (Popov). 19, 21, 26, 28, 29, Sample 8228, east side of Kopalysai; 19, BC 57445, ventral internal mould, x 3; 21, BC 57443, 
ventral internal mould, x 4; 26, BC 57444, ventral internal mould, x 4; 28. BC 57446, dorsal exterior, x 4; 29, BC 57441, internal mould of conjoined 
valves of juvenile specimen, x 4. 20, Sample 8230, Buldukbai-Akchoku section, west side of Kopalysai, BC 57440, latex cast of dorsal interior, x 3. 22— 
25, Sample 818a, Burultas Valley, BC 57442, conjoined valves, dorsal, posterior, lateral and ventral views, x 4. 27. Sample 7613, Akchoku Mountain, 
Kujandysai section, CNIGR 20/1 1989, latex cast of dorsal interior of juvenile, x 3. 

Figs 30, 31 Tesikella necopina (Popov, 1980), Kopalysai, Rukavishnikova. 30, Sample 34, BC 56881, ventral internal mould, x 2. 31. Sample 818a, Burultas 
Valley, BC 57435, x 2.5. 



44 



L.E. POPOV, L.R.M. COCKS AND I.F. NIKITIN 



Shlyginia fragilis (Rukavishnikova, 1956) PI. 6, figs 1 1-25 

1956 Dulankarella fragilis Rukavishnikova: 136, pi. 2, figs 16- 

23. 
1996 Shlyginia fragilis (Rukavishnikova) Nikitin & Popov: 7. 

Holotype. IGNA 28/1369, conjoined valves; Anderken Forma- 
tion, east side of Kopalysai River. 

Material. Eight pairs of conjoined valves, 57 ventral and 53 
dorsal valves from Samples 100b (BC 56989), 620 (BC 56991-99), 
843, 8128a, 8128b, 8135, 8137 (BC 57450), Anderkenyn-Akchoku 
section; Samples 628, 7613, 8258, Kujandysai section; Samples 1 10, 
8230 (BC 57453), 8257, Buldukbai-Akchoku; Sample 8228 (BC 
12881-88, 57447, 48, 52), east side of Kopalysai River; Samples 
390, 818 (BC 57451), 1041a, Burultas Valley; Sample 1018, area 7 
km south-west from Karpkuduk well, Kotnak Mountains, south 
Betpak-Dala. 

Description. Shell concavoconvex, transverse, semielliptical in 
outline, length about two-thirds of the width, with maximum width 
slightly anterior to hinge line or at the hinge line, and 40% as thick as 
long. Cardinal extremities acute to slightly rounded. Anterior com- 
missure rectimarginate. Ventral valve moderately convex in transverse 
profile with maximum thickness at about one-third of valve length. 
Interarea low, planar, apsacline with small triangular delthyrium, 
covered apically by the minute pseudodeltidium. Dorsal valve mod- 
erately convex, slightly geniculate anteriorly with low, planar, 
hypercline interarea and notothyrium covered laterally by disjunct 
chilidial plates. Radial ornament finely parvicostellate with 8-11 
parvicostellae per mm at the anterior margin and 4-8 parvicostellae 
between the accentuated costellae which originate in the umbonal 
area, near the mid-valve and anterior to mid-valve in full grown 
specimens. 

Ventral valve interior with small teeth lacking dental plates. 
Muscle field flabellate, on average 80% as long as wide and 40% as 
long as the valve. Diductor scars large, suboval, deeply impressed 
and completely enclosing small lanceolate adductor scars bisected 
by a fine median ridge. Ventral mantle canals saccate with short 
diverging vascula media. Dorsal interior with a trifid cardinal proc- 
ess widely diverging, low and short socket ridges and strong median 
ridge joined anteriorly with the peripheral rim. Dorsal adductor field 
large, subquadrate. 

Discussion. This species differs from the type species Shlyginia 
declivis Nikitin & Popov (1983: 238, pi. 3, figs 1-5) in its larger size 
and moderately concave lateral profile of the dorsal valve, which is 
also weakly geniculate anteriorly. 



Subfamily PALAEOSTROPHOMENINAE Cocks & Rong, 1989 
Genus GLYPTAMBONITES Cooper, 1956 

Type species. Glyptambonites musculosus Cooper, 1956, from 
the Oranda Formation (Caradoc) of Virginia, U.S.A. 



Glyptambonites sp. 



PI. 4, fig. 6 



Material. Three ventral valves, from Sample 628 (BC 56510), 
Kujandysai, Sample 100, Anderkenyn-Akchoku section. 

DISCUSSION. Glyptambonites is a rare genus in the Anderken 
Formation and also uncommon in the overlying Dulankara Forma- 
tion of the Chu-Ili Range. Its internal features are known from 
specimens from the latter, but not from the Anderken Formation. The 
exterior of our material appears similar to Glyptambonites glyptus 
Cooper, 1956 from the Llandeilo to early Caradoc of Virginia and 
Alabama. 



Genus TESIKELLA gen. nov. 

ETYMOLOGY. After the River Tesik. 

Type SPECIES. Palaeostrophomena necopina Popov, 1980, from 
the Anderken Formation, Chu-Ili Range. 

Diagnosis. Shell profile resupinate, ventral valve with low interarea; 
chilidial plates disjunct; radial ornament unequally parvicostellate; 
ventral interior with double teeth lacking dental plates; ventral muscle 
field enclosed by bilobed bounding ridges: adductor scars short; 
ventral subperipheral rim variably developed; dorsal interior with 
strong median septum coalescing anteriorly with platform. 

DISCUSSION. The subfamily Palaeostrophomeninae has the genera 
Palaeostrophomena, Apatomorpha, Glyptambonites , Ishimia, 
Lepidomena, Titanambonites and Toquimia definitely attributed to it, 
and Goniotrema is possibly a member (Cocks & Rong 2000). Of these, 
all are of normal convexity apart from Palaeostrophomena which is 
generally resupinate and Toquimia, in which resupination develops 
anteriorly in larger specimens. Tesikella is also resupinate and in 
external features resembles Palaeostrophomena apart from the irregu- 
larrugae, which are variable in the latter. However, internally Tesikella 
has a dorsal platform which is absent in Palaeostrophomena, and also 
has a variably developed subperipheral rim in the ventral valve which 
in some specimens is fully developed but in others consists only of 
semi-continuous papillae. The ventral muscle field is enclosed by 
bilobed bounding ridges, which are not present in other members of 
the subfamily, but are developed in other Leptellinidae. 



PLATE 6 

Figs 1-6 Tesikella necopina (Popov, 1980), Sample 8129, Anderkenyn-Akchoku section. 1, BC 57433, ventral internal mould, x 2. 2, BC 57434. ventral 
exterior, x 2. 3, 4, BC 57604, latex cast, x 3, and dorsal internal mould, x 2. 5, 6, BC 57432, latex cast, x 3. and ventral internal mould, x 2. 

Figs 7-10 Sowerbyella (Sowerbyella) rukavishnikovae Popov. 7, Sample 100b, Anderkenyn-Akchoku section. CNIGR 41/1 1522. latex cast of dorsal 
exterior, x 5. 8, Sample 1 10, Buldukbai-Akchoku section, BC 57801, ventral internal mould, x 4. 9, Sample 1018a, 7 km southwest of Karpkuduk well, 
Kotnak Mountains, south Betpak-Dala, CNIGR 44/1 1522, dorsal interior, x 2.5. 10, Sample 100b, CNIGR 38/1 1989, dorsal interior, x 2.5. 

Figs 11-25 Shlyginia fragilis (Rukavishnikova). 11, 12, 17-19, 21, 23, 24, Sample 8228, east side of Kopalysai; 11, BC 57448. ventral internal mould, x 
2; 12, BC 12882, ventral internal mould, x 3; 17-19, BC 57447, ventral, dorsal and lateral views of conjoined valves, x 2; 21, 23, 24, BC 57452, 
conjoined valves, lateral, dorsal and ventral views, x 2. 13-16. Sample 818, Burultas Valley. BC 57451, conjoined valves, ventral, dorsal and lateral 
views, x 4; posterior view of interareas, x 6. 20, Sample 8230, BC 57453, dorsal internal mould, x 3. 22, Sample 8137, Anderkenyn-Akchoku section, BC 
57450, ventral internal mould, x 3. 25, Sample 620, Anderkenyn-Akchoku section, BC 57449, ventral internal mould, x 3. 

Figs 26-33 Sortanella aff. quinquecostata Nikitin & Popov. 26-30, Sample 2538, Akchoku Mountain, Kujandysai section; 26, BC 57488, ventral internal 
mould, x 2; 27-29, BC56771. conjoined valves, dorsal, ventral and lateral views, x 4; 30. BC 57460, ventral internal mould, x 4. 31, Sample 626, 
Anderkenyn-Akchoku section, BC 57458. dorsal exterior, x 2. 32, 33, Sample 100, Anderkenyn-Akchoku section, BC 57459, conjoined valves, dorsal 
and ventral views, x 2. 



UPPER ORDOVICIAN BRACHIOPODS FROM KAZAKHSTAN 



45 














11 






A 






21 




22 













15 



19 




16 





"«S! 




32 



29 



30 



28 




31 




46 



L.E. POPOV, L.R.M. COCKS AND I.F. NIKITIN 



Tesikella necopina (Popov, 1980) 

PI. 5, figs 30, 31, PI. 6, figs 1-6 

1980 Palaeostrophomena necopina Popov: 145, pi. 1, figs 8-1 1 . 

HOLOTYPE. CNIGR 15/1 1523 (L=l 1.4, W=16.8), dorsal internal 
mould, Anderken Formation, east side of Kopalysai, Sample 127/K- 
1970. 

Material. Five pairs of conjoined valves, 60 ventral and 43 dorsal 
valves from Samples 8128, 8129 (BC 57432-34, 57604), 8138, 
Anderkenyn-Akchoku section; Sample 7613, Kujandysai section; 
Sample 127/K-1970 and Rukavishnikova (1956) Sample 34 (BC 
56881), east side of Kopalysai; Sample 818a (BC 57435), Burultas 
Valley; Sample 1024b, east side of Karatal near Sorbulak well; 
Samples 1018, 1018a, area 7 km south-west of Karpkuduk well, 
Kotnak Mountains. 

DESCRIPTION. Shell profile resupinate, transversely subrectangular 
outline, about 55-60% as long as wide with maximum width at the 
hinge line. Cardinal extremities slightly acute to near right angled. 
Anterior commissure rectimarginate, broadly rounded. Ventral valve 
with lateral profile slightly convex in the umbonal area, gently 
concave anteriorly. Ventral interarea low, planar, catacline with a 
well developed, narrow pseudodeltidium. Dorsal valve with moder- 
ately convex lateral profile, flattened posteriorly with low anacline 
interarea and separate chilidial plates. Radial ornament parvicostellate 
with 10-12 parvicostellae per mm in mature specimens and accentu- 
ated costellae of two-three generations. 

Ventral interior with strong, double teeth lacking dental plates and 
large divided musle field with strong diductor scars and muscle 
bounding ridges extending anteriorly to the mid-valve. Adductor 
scars small, strip-like, divided by a fine median ridge, about half the 
length of the diductor scars. Subperipheral rim variably developed, 
posterior to which is a weak median ridge. Mantle canals saccate 
with very short vascula media branching just beneath the anterior 
margin of the diductor scars. Dorsal interior with trifid cardinal 
process on a low notothyrial platform and low, widely diverging 
socket ridges. Median septum strong and narrow, about 75% as long 
as the valve and joined anteriorly to a low subperipheral rim. 
Adductor scars radially arranged with smaller anterior pair extend- 
ing anteriorly to mid-valve. 

Discussion. Popov (1980) originally attributed the species to 
Palaeostrophomena, but since then the internal characteristics, par- 
ticularly of the ventral valve, have become known, and it is clear that 
this species cannot properly be attributed to that genus. 



DISCUSSION. These specimens closely resemble Sortanella 
quinquecostata Nikitin & Popov (1996: 9) in radial ornament, 
posteriorly subcarinate ventral valve and gently uniplicate anterior 
commissure. The type locality is about 400 km to the north-west of 
our localities, and occurs in the overlying Dulankara Regional Stage 
which is late Caradoc rather than early to middle Caradoc. Because 
no dorsal interiors are known, the species cannot yet be identified 
from the Anderken Formation with certainty. 



Subfamily AEGIROMENINAE Havh'ek, 1961 
Genus CHONETOIDEA Jones, 1928 



Chonetoidea sp. 



PI. 5, figs 4, 5 



Type SPECIES. Plectambonites papillosa Reed, 1905, from the 
Slade and Redhill Mudstone Formation (Middle Ashgill), 
Pembrokeshire, Wales. 

Material. One internal and one external mould of a pair of 
conjoined valves, BC 56537 (L=3.2, W=5.1) from Sample 8255, 
Anderkenyn-Akchoku section. 

Description. Shell planoconvex, transverse, semielliptical in out- 
line with maximum width at the hinge line. Cardinal extremities 
acute. Anterior commissure rectimarginate. Ventral valve gently 
convex in lateral profile with maximum thickness slightly anterior to 
the apex. Interarea low, planar, anacline with minute apical 
pseudodeltidium. Dorsal valve flat with a low, anacline interarea. 
Chilidial plates separate. Radial ornament finely and unequally 
parvicostellate with five accentuated primary ribs and four second- 
ary costellae originating at about mid valve length. 

Ventral interior with small, bilobate muscle field bisected 
posteriorly by low median ridge. Dorsal interior with simple under- 
cut cardinal process joined to minute socket ridges subparallel to the 
hinge line. Median ridge originating anterior to the deep alveolus and 
extending to mid- valve. Six septulae lateral to the anterior part of the 
median ridge near the mid-valve. 

DISCUSSION. These specimens resemble Chonetoidea virginica 
Cooper ( 1956: 805 ), from the Edinburg Formation of Virginia, in the 
size and outline of the planoconvex shell, unequally parvicostellate 
radial ornament, and number and arrangement of septulae in the 
dorsal valve. 



Family HESPERONOMIIDAE Cooper, 1956 
Genus ANOPTAMBON1TES Williams, 1962 



Family XENAMBONITIDAE Cooper, 1956 

Subfamily XENAMBONITINAE Cooper, 1956 

Genus SORTANELLA Nikitin & Popov, 1996 

TYPE SPECIES. Sortanella quinquecostata Nikitin & Popov, 1 996, 
from the Dulankara Regional Stage (Upper Caradoc), north Betpak- 
Dala, Kazakhstan. 

Sortanella aff. quinquecostata Nikitin & Popov, 1996 

PI. 6, figs 26-33 

Material. Ten pairs of conjoined valves, 7 ventral and 8 dorsal 
valves from Samples 100 (=K98/1970) (BC 57459), 626 (BC 57000, 
57006-9, 57458), Anderkenyn-Akchoku section; Samples 628, 2538 
(BC 5677 1,568 10, 570 10-03, 57460, 88), Kujandysai section; Tesik 
River, Sample 948 (BC 57002-5). 



TYPE SPECIES. Leptaena grayae Davidson, 1883, from the 
Craighead Limestone (Upper Caradoc) of Girvan, Scotland. 

Anoptambonites convexus sp. nov. 

PI. 7, figs 5-26; Figs 12.1-12.6 

1986 Anoptambonites sp.; Kolobova & Popov, pi. 1, figs 7. 8. 

ETYMOLOGY. After convexus, Latin - convex. 

HOLOTYPE. BC 57462, PI. 7, figs 5, 6, a dorsal interior from 
Sample 100, Akchoku Mountain. 

Material. 36 pairs of conjoined valves, 21 ventral and 14 dorsal 
valves from Samples 1 00 (=K98/1 970) (BC 56540, 570 1 4-32, 57462, 
64, 69, 71), 626 (BC 57466, 68), 8214 (BC 57480), 8223 (BC 
57063-68), 8223b (BC 57467), Anderkenyn-Akchoku section; Sam- 
ples 628 (BC 56530, 42, 57043-51). 2538 (BC 56531, 57052-61, 



UPPER ORDOVICIAN BRACHIOPODS FROM KAZAKHSTAN 



47 



Table 15 Measurements of complete shells and ventral valves of 
Anoptambonites convexa sp. nov., Acculina-Dularikarella Association, 
samples 100 and 626 from Anderkenyn-Akchoku section. 





Lv 


W 


T 


Lv/W 


T/Lv 


N 


18 


17 


8 


17 


8 


X 


12.1 


16.5 


6.9 


71.6% 


47.1% 


s 


4.24 


5.26 


1.86 


7.5 


6.3 


MIN 


5.8 


8.2 


3.5 


57.8% 


39.4% 


MAX 


18.0 


23.9 


9.6 


86.5% 


58.9% 



Table 16 Measurements of complete shells and ventral valves of 

Anoptambonites convexa sp. nov.. Parastrophina-Kelterella Association, 
samples 2538, 8217 and 8256 from Kujandysai section. 





Lv 


W 


T 


Lv/W 


T/Lv 


N 


11 


11 


4 


11 


4 


X 


6.4 


9.9 


2.8 


65.5% 


44.8% 


s 


1.13 


2.33 


0.57 


7.6 


6.1 


MIN 


5.2 


6.6 


2.0 


57.0% 


38.5% 


MAX 


9.0 


15.3 


3.2 


79.3% 


51.6% 



57461, 63, 65), 8217, 8256 (BC 57070-73), Kujandysai section; 
Samples 8230 (BC 57069), 8231-40, Buldukbai-Akchoku. 

DESCRIPTION. Shell concavoconvex, transverse, semielliptical in 
outline, about 72% as long as wide with maximum width at hinge 
line and thickness 47% of valve length. Cardinal extremities slightly 
acute to rectangular. Anterior commissure rectimarginate. Ventral 
valve carinate posteriorly, strongly convex in lateral profile with the 
maximum thickness at the point of geniculation somewhat anterior 
to mid-length. Beak pointed and slightly erect posterior to the hinge 



line. Ventral interarea steeply apsacline to procline with mainly open 
delthyrium covered apically by the minute pseudodeltidium. Dorsal 
valve gently and unevenly concave in lateral profile, flat to mid- 
length. A shallow sulcus originates at the umbo and fades towards the 
anterior margin. Dorsal interarea anacline with convex chilidium. 
Radial ornament finely and near equally multicostellate with 28^-3 
primary ribs originating near the umbo and 4—6 ribs per mm at the 
anterior margin. Concentric ornament of fine, evenly spaced fila. 

Ventral valve with teeth lacking dental plates. Muscle field small, 
cordate, bisected by fine median ridge separating small, lanceolate 
adductor scars. Vascula media short, widely diverging. Dorsal inte- 
rior with undercut cardinal process bearing up to 8 ridges on both 
sides of strong central lobe. Lophophore platform semielliptical, 
bordered by a high rim joined to the median septum. Dorsal adductor 
muscle field subrectangular, about one-third as long as the valve. 

VARIABILITY. The average size of Anoptambonites convexus from 
the Acculina-Dulankarella Association, which typically occurs 
within the nodular limestone deposited on the flanks of carbonate 
mud mounds (Sample 100), is one and half to two times larger than 
the average size of the shells from the pockets in the mud mound core 
(Samples 2538 and 8231-40) and the overlying bedded limestone 
(Samples 628, 8217, 8223, 8256) in which the Parastrophina- 
Kellereila Association characteristically occurs. However, the 
specimens from the Parastrophina-KellereUa Association retain a 
similar outline, transverse and lateral profile of the shells from the 
Acculina-Dulankarella Association and are also characterized by 
their multicostellate ornament which consists of 26 to 36 primary 
ribs and 4 to 6 ribs along the anterior margin of full grown specimens. 

DISCUSSION. This species differs from Anoptambonites grayae 
(Davidson), as revised by Williams (1962: 1 7 1 ) from the Craighead 




Fig. 12 1-6. Anoptambonites convexus sp. nov. 1, 4, 5, Anderkenyn-Akchoku section, 1, Sample 8223b, BC 57467, ventral exterior, x 3; 4, 5, Sample 100. 
4, BC 57470, ventral internal mould, x 2; 5, BC 57472. ventral exterior, x 2. 3, 6, Buldukbai section, 3, Sample 8230, BC 57069, ventral internal mould, 
x 3, 6, Sample 8231, BC 57473, ventral internal mould, x 1.5. 2. Kujandysai section, Sample 8256, ventral exterior, x 2. 

7-14, Sowerbyella (Sowerbyella) aff. ampla (Nikitin & Popov), 7-13, Kujandysai section, Sample 2538, 7, BC 57082, ventral exterior, x 3; 8,9, BC 
57482, lateral and ventral views of ventral exterior, x 4; 10, BC 57081, ventral internal mould, x 3; 11, 12, BC 57479, anterior and ventral views of 
ventral exterior, x 3; 13, BC 57481, ventral exterior, x 3. 14, Anderkenyn-Akchoku section. Sample 8214, BC 57480. dorsal exterior, x 2.5. 



48 



L.E. POPOV, L.R.M. COCKS AND I.F. NIKITIN 



Limestone ( Upper Caradoc) of Girvan, in having a strongly concavo- 
convex lateral profile, a cardinal process with up to 8 vertical ridges 
on the lateral lobes, a shorter median septum and a relatively small 
lophophore platform extending anteriorly only to the mid-valve. 
Anoptambonites convexus differs from two somewhat younger 
Kazakh species, A. subcarinatus Nikitin & Popov (1996:10) from 
the north Betpak-Dala and A. kovalevskii Popov, Nikitin & Cocks 
(2000), from the Dulankara Mountains, both from the Dulankara 
Regional Stage (Upper Caradoc to lowermost Ashgill), in having 
equally multicostellate radial ornament and a weakly geniculate 
ventral valve profile, with the maximum height anterior to the mid- 
valve and near the point of geniculation. It also differs from the 
former species in having a rectimarginate anterior commissure. 

Anoptambonites orientalis Popov, 1980 PI. 7, figs 1-4, 27 

1980 Anoptambonites orientalis Popov: 149; pi. 2, figs 12-17. 

Holotype. CNIGR 30/11523, dorsal internal mould, from the 
Anderken Formation, Anderkenyn-Akchoku section, Sample 100b. 

Material. Two pairs of conjoined valves, 14 ventral and 26 dorsal 
valves from Samples 100b, 8128a (BC 56533, 57476), 8128b, 8137 
(BC 57478), Anderkenyn-Akchoku section; Sample 8230, 
Buldukbai-Akchoku section, and Sample 1018, about 7 km south- 
west of Karpkuduk well, Kotnak Mountains. 

DISCUSSION. Detailed description of this species was provided by 
Popov (1980). It differs from the contemporaneous Anoptambonites 
convexus sp. nov. as well as from the somewhat younger A. 
subcarinatus Nikitin & Popov, 1996 and A. kovalevskii Popov, 
Nikitin & Cocks, 2000 in having a flattened shell, a carinate ventral 
valve with gently and evenly convex lateral profile lacking 
geniculation, a very weakly concave dorsal valve, and in the pres- 
ence of a small but well-defined pseudodeltidium. The platform in 
mature specimens of A. orientalis reaches up to 75% of the valve 
length, whereas in A. convexus and A. kovalevskii it barely exceeds 
half the valve length. 



Genus KASSINELLA (KASSINELLA) Borissiak, 1956 

Type species. Kassinella globosa Borissiak, 1 956, from the Lower 
Ashgill Kulunbulak Formation, Tarbagatai Range, Kazakhstan. 



Kassinella (Kassinella)? sp. 



PI. 3, fig. 24 



Material. One dorsal internal mould, BC 56497, from Sample 
2531, Anderkenyn-Akchoku section. 

Discussion. A single dorsal valve shows the undercut cardinal 
process and lack of both bema and side septa characteristic of the 
Hesperomenidae. It possesses an evenly semicircular platform and a 
median septum not extending anterior of the platform. This evenly 



semicircular platform is unlike most species of Hesperomena, 
Anoptambonites, Aulie and Chaganella: within the family only 
Kassinella (Kassinella) has such a platform. However, without 
knowledge of the ventral interior and the exterior of both valves, 
generic assignment can only be provisional. Kassinella (Kassinella) 
is known from the late Caradoc and Ashgill of Kazakhstan, South 
China. Australia, Scotland, Bohemia and Sweden (Zhan & Cocks 
1998:36), but if the record from the Anderken Formation is con- 
firmed then this may be its earliest occurrence. 



Family SOWERBYELLIDAE Opik, 1930 

Subfamily SOWERBYELLINAE Opik, 1930 

Genus SOWERBYELLA (SOWERBYELLA) Jones, 1928 

Type SPECIES. Leptaena sericea J. de C. Sowerby, 1 839, from the 
Horderley Sandstone (Lower Caradoc) of Shropshire, England. 

Sowerby ella (Sowerby ella) rukavishnikovae Popov, 1980 

PI. 6, figs 7-10 

1980 Sowerbyella rukavishnikovae Popov: 151, pi. 2, figs 1-4. 
1984 Sowerbyella rukavishnikovae Popov; Nikitin & Popov in 
Kleninaero/.:150. pi. 16. figs 17-22. 

HOLOTYPE. CNIGR 40/1 1523, a dorsal internal mould from the 
Anderken Formation, Anderkenyn-Akchoku section, Sample 100b. 

Material. 70 ventral and 133 dorsal valves. Samples 100b (BC 
56554-5), 848, 8128a, 8128b, 8137, Anderkenyn-Akchoku section; 
Sample 7613, Kujandysai section; Samples 1 10 (BC 57801), 8229, 
8230, 8257. Buldukbai-Akchoku; Sample 8228, east side of the 
Kopalysai River; Sample 1024b, east side of Karatal Valley, near 
Sorbulak well; Samples 1018, 1018a, 7 km south-west of Karpkuduk 
well, Kotnak Mountains. 

Discussion. Description, discussion and basic statistics of this 
species were provided by Popov (1980). Some further specimens 
from the Anderken Formation are illustrated here. This species is 
also reported from the upper Bestamak and lower Sargaldak Forma- 
tions of the Chingiz Range (Nikitin & Popov in Klenina et al. 1984). 

Sowerbyella (Sowerbyella) aff. ampla (Nikitin & Popov, 
1996) Figs 12.7-12.14 

1986 Anisopleurella sp. Kolobova & Popov: pi. 1, fig. 6. 
aff. 1996 Anisopleurella ampla Nikitin & Popov: 12, figs 5K-R. 

Material. 14 pairs of conjoined valves. 25 ventral and 8 dorsal 
valves from Samples 100 (=K98/70), 626, 2531, 8214 (BC 57480), 
8215, 8223a, 8223b, Anderkenyn-Akchoku section; Samples 628, 
2538 (BC 57081, 82, 57479, 81, 82), Kujandysai section; Sample 
948 (BC 57084-85), Tesik River. 



PLATE 7 

Figs 1-4, 27 Anoptambonites orientalis Popov. 1-3, Sample 8 1 28a. Anderkenyn-Akchoku section; 1, BC 56533, latex cast of dorsal interior, x 4; 2, 3, BC 
57476. ventral internal mould and latex cast, x 2. 4, 27, Sample 8137. Anderkenyn-Akchoku section. BC 57478. latex cast and dorsal internal mould of 
immature specimen, x 4. 

Figs 5-26 Anoptambonites convexus sp. nov. 5, 6. 9, 10, 18, 20-22, Sample 100, Anderkenyn-Akchoku section; 5, 6, BC 57462. holotype, dorsal internal 
mould and latex cast, x 2; 9, 10, BC 57464, posterior view of conjoined valves showing interareas, x 1.5 and x 5; 18, BC 57469. ventral exterior, x 2; 20- 
22, BC 57471, conjoined valves, ventral, lateral and dorsal views, x 2. 7, 8, 11-16, Sample 2538, Akchoku Mountain, Kujandysai section; 7, BC 57461, 
ventral internal mould, x 3; 8, CNIGR 7/12361, ventral exterior, x 3; 11, 12, BC 57465, latex cast and dorsal internal mould, x 3; 13-16, BC 57463, 
conjoined valves, dorsal, lateral, ventral and posterior views, x 3. 17, 23-26. Sample 626. Anderkenyn-Akchoku section; 17, BC 57466. ventral internal 
mould, x 1.5; 23-26, BC 57468, conjoined valves, dorsal, posterior, lateral and ventral views, x 2. 19, Sample 628 (=K- 107/70), west side of Kujandysai, 
BC 56542, ventral internal mould, x 2. 



UPPER ORDOVICIAN BRACHIOPODS FROM KAZAKHSTAN 



49 




50 



L.E. POPOV, L.R.M. COCKS AND I.F. NIKITIN 



Description. Shell concavoconvex, transverse, semielliptical in 
outline, about 56% as long as wide, with maximum width at hinge 
line. Cardinal extremities acute and slightly alate. Anterior commis- 
sure broadly rounded and rectimarginate. Ventral valve moderately 
and unevenly convex in profile with the maximum thickness at about 
quarter valve length. Ventral interarea strongly apsacline and slightly 
curved in cross-section, with small triangular delthyrium covered 
apically by pseudodeltidium. Dorsal valve moderately concave, with 
hypercline interarea; notothyrium covered laterally by chilidial plates. 
Radial ornament unequally and finely parvicostellate with closely- 
spaced parvicostellae varying from 9 to 1 2 per mm. Five accentuated 
ribs originating at the umbo and four accentuated costellae inclined 
between them in the mid-valve. Indistinct radial plications devel- 
oped sometimes along the accentuated ribs. About 6-9 strong 
concentric rugellae inclined below 30-40° towards the hinge line. 

Ventral interior with small teeth lacking dental plates and small 
bilobate muscle field with flabellate diductor scars completely sur- 
rounding small lanceolate adductor scars. Dorsal interior with 
undercut cardinal process, small curved socket ridges, bilobed bema, 
short median ridge and two widely divergent side septa. 

Discussion. The Anderken specimens strongly resemble 
Anisopleurella ampla Nikitin & Popov (1996) from the slightly 
younger Dulankara Regional Stage of north Betpak-Dala, Central 
Kazakhstan, in size and general shell shape, but differ slightly in 
having a finer radial ornament with 9-12 parvicostellae per mm 
instead of 7-10 in the Dulankara specimens, and an uneven ventral 
valve lateral profile. Anisopleurella ampla is reassigned here to 
Sowerbyella (Sowerbyella) mainly because it has two pairs of side 
septa including a closely-spaced more prominent pair. In the variable 
development of strong concentric ornament it is similar to the 
subgenus S. (Rugosowerbyella), but differs in having a median 
septum and the two pairs of side septa, and in the small, weakly 
impressed ventral muscle field. 



Genus ANISOPLEURELLA Cooper, 1956 

TYPE SPECIES. Anisopleurella tricostellata Cooper, 1 956, from the 
Pratt Ferry Formation (Llandeilo) of Alabama, U.S.A. 

DISCUSSION. The relationships of Anisopleurella and other genera 
are unresolved. The Ordovician genera within the subfamily are 
Sowerbyella, Anisopleurella, Eochonetes, Eoplectodonta, 
Gunningblandella and our new genera Olgambonites and 
Zhilgyzambonites. Eochonetes and Eoplectodonta may be inter- 
preted as Sowerbyella with hinge lines modified to form denticles 
(which resulted in the subsequent loss of dental plates), and 



Gunningblandella and Olgambonites are resupinate modifications 
of Sowerbyella, which was a very plastic stock. Anisopleurella has a 
much more distinctive and erect dorsal median septum and side septa 
than Sowerbyella, with the side septa bisecting the divided bema but 
not reaching to the anterior edge of it. Both genera occur in rocks of 
Llandeilo age, but which of the two was ancestral is not known. 



Anisopleurella sp. 



PI. 8, figs 1,2,5 



Material. One ventral and one dorsal internal mould from Sam- 
ples 2531 (BC 57489), 8255 (BC 56539), Anderkenyn-Akchoku 
section. 

Discussion. These specimens can be firmly assigned to 
Anisopleurella because they have the characteristic parvicostellate 
ornament with three accentuated ribs, a dorsal median ridge and a 
pair of widely diverging prominent side septa bisecting a bilobed 
bema. They are somewhat comparable to, and may be conspecific 
with, Anisopleurella yichangensis Zeng, 1 987 from the early Caradoc 
Miaopo Formation of Hubei. South China, but the absence of well- 
preserved exteriors of the Chinese species makes detailed comparison 
impossible. 

Genus OLGAMBONITES gen. nov. 

Etymology. After the late Olga Ivanova Nikiforova, a pioneer in 
brachiopod studies. 

TYPE SPECIES. Olgambonites insolita sp. nov. from the Anderken 
Formation. Chu-Ili Range. 

DIAGNOSIS. Shell convexiconcave; anterior commissure 
rectimarginate to weakly uniplicate; ventral interarea procline to 
slightly apsacline with apical pseudodeltidium; dorsal interarea 
anacline with separate chilidial plates; radial ornament unequally 
parvicostellate; ventral interior with small teeth lacking dental plates 
and small bilobed muscle field with short adductor scars completely 
separating larger diductor scars; ventral mantle canals lemniscate; 
dorsal interior with simple undercut cardinal process joined to 
narrow socket ridges; fine median ridge and bilobed bema bordered 
by rim and crossed by up to 8 side septa. 

Discussion. Olgambonites possesses an undercut cardinal proc- 
ess, divided bema and side septa in the dorsal valve; all characteristic 
of the Sowerbyellidae, but it differs from most genera in the family 
in having a convexiconcave shell. The only other resupinate genus is 
Gunningblandella (Percival 1979b) from the Caradoc of Australia, 
but Olgambonites differs from that genus in having a dorsal median 
septum and numerous side septa. 



PLATE 8 

Figs 1, 2, 5 Anisopleurella sp. 1, 2, Sample 8255, Anderkenyn-Akchoku section, BC 56539, dorsal internal mould and latex cast, x 12. 5, Sample 2531. 

Anderkenyn-Akchoku section, BC 57489, ventral internal mould, x 7. 
Figs 3, 4, 6-10, 12, 13 Zhilgyzambonites extenuata gen. et sp. nov. Anderkenyn-Akchoku section. 3, 9, 10, 12, 13, Sample 8255; 3, BC 57493, ventral 

internal mould, x 6; 9, BC 56538, latex cast of conjoined ventral and dorsal interiors, x 4; 10, BC 57494, ventral internal mould, x 6; 12, 13, BC 12915, 

holotype, latex cast and internal mould of dorsal interior, x 5.5. 4, 7, 8, Sample 253 1 ; 4, BC 57490. latex cast of ventral exterior, x 6: 7, 8, BC 57492, 

dorsal internal mould and latex cast, x 5. 6, Sample 8215, BC 57491. latex cast of dorsal exterior, x 6. 
Figs 11, 14-20 Olgambonites insolita gen. et sp. nov. Sample 8255, Anderkenyn-Akchoku section. 11, BC 56535. dorsal external mould, x 4. 14, 15, BC 

56534. latex cast and ventral internal mould, x 4. 16, 17, BC 56664, latex cast and ventral internal mould, x 4. 18, BC 57592. ventral exterior, x 4. 19, 20, 

BC 56663. holotype, internal mould and latex cast of dorsal valve, x 4. 
Figs 21-33 Gacella institata sp. nov. 21-25, 32, 33. Sample 100. Anderkenyn-Akchoku section; 21-25, BC 57496, conjoined valves, posterior, anterior, 

dorsal, ventral and lateral views, x 2; 32, 33, BC 57500. conjoined valves, holotype, dorsal and ventral internal moulds, x 2. 26, 27. Sample 85258, 

Kujandysai section, BC 56576, latex cast and dorsal internal mould, x 3. 28-31. Sample 626, Anderkenyn-Akchoku section, BC 57497, conjoined valves, 

anterior, ventral, lateral and dorsal views, x 2. 



UPPER ORDOVICIAN BRACHIOPODS FROM KAZAKHSTAN 



51 




52 



L.E. POPOV, L.R.M. COCKS AND I.F. NIKITIN 



Olgambonites insolita sp. nov. PI. 8. figs 1 1, 14-20 

Etymology. After insolitus, Latin -extraordinary. 

Holotype. BC 56663, a dorsal valve (PL 8, figs 19, 20) from the 
Anderken Formation, Sample 8255, Anderkenyn-Akchoku section. 

Material. 3 ventral (BC 56534-36, BC 56664) and 2 dorsal 
valves (BC 56538, BC 56663) from the type locality. 

Description. Shell convexoconcave, semielliptical in outline, 
about half as long as wide, with maximum width at the hinge line, 
and acute cardinal extremities. Ventral valve gently concave in 
lateral profile with apsacline interarea and narrow convex 
pseudodeltidium. Dorsal valve gently convex with anacline 
pseudointerarea. Radial ornament inequally parvicostellate with 7- 
9 primary accentuated ribs and two generations of accentuated 
costellae. Very fine parvicostellae, about 10-1 1 per mm. Concentric 
ornament of fine evenly spaced rugellae and very fine crowded 
growth lamellae in the anterior half of the shell. 

Ventral interior with small teeth and highly raised, rounded, 
subrectangular muscle field bordered anteriorly by the steep rim. 
Dorsal interior with simple undercut cardinal process joined to the 
socket ridges and small alveolus. Bema entire, highly raised anteriorly 
and bordered by a steep rim. Short median ridge originating anterior 
to bema and joined to the subperipheral rim. 

Discussion. The new species is known only from the type locality. 



Genus ZHILGYZAMBONITES gen. nov. 
Etymology. After Zhilgyz well, Betpak-Dala Desert. 

Type species. Zhilgyzambonites e.xtenuata sp. nov. from the 
Anderken Formation, Chu-Ili Range. 

Diagnosis. Shell concavoconvex, with rectimarginate posterior 
commissure, ventral interarea apsacline with delthyrium completely 
covered by pseudodeltidium; dorsal interarea anacline with com- 
plete chilidium; radial ornament finely and unequally parvicostellate; 
ventral valve with small teeth lacking dental plates; ventral muscle 
field small, highly raised anteriorly; dorsal interior with undercut 
cardinal process joined to socket ridges, deep alveolus and strongly 
elevated, entire bema; median ridge fine, originating anteriorly to 
bema and joined anteriorly to the subperipheral rim. 

DISCUSSION. Zhilgyzambonites is somewhat similar to Aulie 
(Nikitin & Popov in Klenina et al. 1984). which is within the Hes- 
peromenidae, in the characteristic pseudodeltidium and chilidium. 
the ventral valve, which lacks dental plates and has a small muscle 
field which is strongly raised anteriorly, and the dorsal interior with 
the deep alveolus and an undercut cardinal process: but it differs in 
having an elevated bema and a short median ridge between the 
anterior margin of the bema and the subperipheral rim. However, the 
presence of a bema in Zhilgyzambonites places it within the 



Table 17 Measurements of ventral valves of Zhilgyzambonites extenuata 
sp. nov., Samples 2531, 8251. 8255, from Anderkenyn-Akchoku section. 



Lv 



W 



LvAV 



N 
X 

s 

MIN 
MAX 



7 

3.7 

0.78 

2.2 

4.4 



7 


7 


5.2 


70.9% 


0.97 


5.4 


3.2 


61.1% 


6.2 


78.6% 



Sowerbyellidae on the criteria established by Cocks & Rong (1989). 
The dorsal interior of the new genus somewhat resembles 
Diambonioidea (Zeng 1987), but the Kazakh genus differs from the 
latter in having a strongly raised ventral muscle field, a ventral 
median ridge anterior to the muscle field, and an undercut cardinal 
process; the simple, not undercut, cardinal process of Diambonioidea 
places it within the Grorudiidae (Cocks & Rong 2000). 

Zhilgyzambonites extenuata sp. nov. 

PI. 8, figs 3, 4, 6-10. 12, 13 

Etymology. After extenuatus, Latin - little, weak. 

Holotype. BC 12915, a dorsal valve (PI. 8, figs 12, 13) from the 
Anderken Formation, Sample 8255, Anderkenyn-Akchoku section. 

Material. 2 pairs of conjoined valves, 10 ventral and 10 dorsal 
valves from Samples 2531 (BC 57490, 92), 8255 (BC 12915-21, 
57493, 94) and possibly 82 1 5 ( BC 57089, 9 1 , 5749 1 ), Anderkenyn- 
Akchoku section. 

Description. Shell concavoconvex, semielliptical in outline, on 
average 70% as long as wide with maximum width at the hinge 
line. Cardinal extremities nearly right angled. Ventral valve mod- 
erately and evenly convex in lateral profile with apsacline interarea 
and narrow convex pseudodeltidium completely covering the 
delthyrium. Dorsal valve moderately concave with linear, anacline 
interarea and convex chilidium. Radial ornament unequally 
parvicostellate with 7-9 accentuated primary ribs and two genera- 
tions of accentuated costellae. Parvicostellae very fine, about 
10-11 per mm. Concentric ornament of fine evenly spaced 
rugellae and very fine crowded growth lamellae in the anterior 
half of the shell. 

Ventral valve interior with small teeth and highly raised, rounded, 
subrectangular muscle field about 20% valve length and bordered 
anteriorly by a steep rim. Dorsal interior with simple undercut 
cardinal process joined to narrow, strongly curved socket ridges; 
small alveolus. Dorsal adductor scars with anterior pair slightly 
larger than posterior, strongly impressed, divided by a pair of 
transmuscle septa. Bema entire, strongly raised anteriorly and bor- 
dered by a steep rim. Short median ridge originating anteriorly to 
bema and joined to the subperipheral rim. 

Discussion. As far as is yet known, this is the only species within 
the new genus. The material from Sample 8215 does not include 
valve internals and is thus only doubtfully referred to the species. 



Table 18 Measurements of dorsal valves of Zhilgyzambonites extenuata sp. nov.. Samples 253 1 . 825 1 . 8255. from Anderkenyn-Akchoku section. 





Ld 


W 


Ml 


Mw 


LP1 


LPw 


BB1 


BBw 


LdAV 


Ml/Ld 


Ml/Mw 


PLl/Ld 


LPwAV 


BBwAV 


N 


8 


8 


3 


3 


3 


3 


3 


3 


8 


3 


3 


3 


3 


3 


X 


3.8 


6.1 


1.5 


2.6 


2.9 


4.8 


0.3 


2.3 


62.5% 


35.1% 


58.6% 


67.4% 


73.9% 


35.7% 


s 


0.53 


0.67 


0.17 


0.40 


0.21 


0.20 


0.06 


0.72 


6.0 


2.4 


10.3 


7.2 


0.3 


12.6 


MIN 


3 


5.2 


1.4 


2.2 


2.7 


4.6 


0.3 


1.7 


54.0% 


33.3% 


46.7% 


62.2% 


73.5% 


26.2% 


MAX 


4.5 


6.8 


1.7 


3 


3.1 


5 


0.4 


3.1 


71.7% 


37.8% 


65.4% 


75.6% 


74.2% 


50.0%Order 



UPPER ORDOVICIAN BRACHIOPODS FROM KAZAKHSTAN 



53 



ORTHOTETIDA Waagen, 1884 

Suborder ORTHOTETIDINA Waagen, 1884 

Superfamily CHILIDIOPSOIDEA Boucot, 1959 

Family CHILIDIOPSIDAE Boucot. 1959 

Subfamily GACELLINAE Williams & Brunton, 2000 

Genus GACELLA Williams, 1962 

TYPE SPECIES. Gacella insolita Williams, 1962, from the Stinchar 
Limestone (Lower Caradoc), Girvan, Scotland. 

Gacella institata sp. nov. PI. 8, figs 21-33, PI. 9, figs 1-4 

Etymology. After instita, Latin - a swathe. 

HOLOTYPE. BC 57500, PI. 8, figs 32, 33, from the Anderken 
Formation, Sample 100, Anderkenyn-Akchoku section. 

Material. 10 pairs of conjoined valves, 2 ventral and 8 dorsal 
valves, from Samples 100 (=K-98/1970) (BC 57496, 98. 99, 57500), 
626 (BC 57497), Anderkenyn-Akchoku section; Samples 628, 85258 
(BC 57092-3, 56576), Kujandysai Section.. 

DESCRIPTION. Shell subequally biconvex, transverse, semielliptical 
in outline about 80% as long as wide, with maximum width at hinge 
line. Cardinal extremities acute to rectangular. Anterior commissure 
gently uniplicate. Ventral valve convex in lateral profile with maximum 
thickness slightly anteriorto the umbo and with flattened sides. Sulcus 
shallow, originating about 2-3 mm from the umbo. Ventral interarea 
planar, apsacline with a broad, convex pseudodeltidium perforated 
apically by a minute foramen. Lateral profile of the dorsal valve gently 
convex with maximum thickness near the anterior margin. Dorsal 
median fold low, rounded in cross-section, originating in the umbonal 
area but very weakly defined until the mid-valve. Dorsal interarea 
planar, anacline, with convex chilidium. Radial ornament unequally 
parvicostellate with two to three generations of acentuated ribs, about 
4-5 per 3 mm along the anterior margin of mature specimens. 
Concentric ornament of numerous fine growth lamellae anteriorly. 

Ventral interior with teeth supported by long subparallel but 
slightly divergent dental plates and narrow, elongate subtriangular 
muscle field divided by median ridge. Numerous fine crenulations 
on the outer surface of the teeth. Dorsal interior with bilobed cardinal 
process on a high notothyrial platform: adductor scars elongated 
slightly, shorter than half valve length, crossed by two pairs of short 
transmuscle septa. Median ridge fine and faint, originating some 
distance from the notothyrial platform. 

Measurements. (47 1/12375) conjoined valves, L=l 4.0, W= 1 6.9, 
T=6.3, Sw=8.9; (474/12375) conjoined valves, L=20.4, W=21.0, 
T=11.2, Sw=8.7; (475/12375) conjoined valves, L=22.9, W=24.5, 
T=10.2, Sw=12.8; (476/12375) conjoined valves, L=16.5, W=19.8, 
T=6.5, Sw=11.2; (479/12375) conjoined valves, L=10.5, T=6.7, 
Sw=6.7. 

Discussion. This species differs from others of the genus in 
having a concentric ornament of strong growth lamellae and coarser 
accentuated ribs. It can be compared to Gacella ponderosa Williams, 
1962, from the Confinis Formation (Llandeilo) of Girvan, south 
Scotland, in the general shape and size of the shell, but differs in 
having a less convex lateral profile in both valves in addition to the 
patterns of radial and concentric ornament. 

Gacella institata differs from G. sulcata Misius (in Misius & 
Ushatinskaya 1 977), from the Tabylgaty Formation (Upper Caradoc) 
of the Moldo-Too Range, Kyrgizstan. in having a shallow ventral 
median sulcus and low dorsal median fold usually originating in the 
umbonal region; the convex, not flat, lateral profile of the ventral 
valve, and in the absence of geniculation in the dorsal valve. 



Suborder TRIPLESHDINA Moore, 1952 

Superfamily TRIPLESIOIDEA Schuchert, 1913 

Family TRIPLESIIDAE Schuchert, 1913 

Genus TRIPLESIA Hall, 1859 

Type SPECIES. Atrypa e.xtans Emmons, from the Trenton Group 
(Caradoc), New York. U.S.A. 



Triplesia sp. 



PI. 9, figs 22-26 



Material. Two ventral and three dorsal valves (BC 575 12) from 
Sample 8228. Kopalysai. 

DESCRIPTION. Shell dorsibiconvex, transverse suboval in outline, 
about 80% as long as wide with uniplicate anterior commissure. 
Ventral valve moderately convex, with maximum thickness at about 
one-third valve length. Ventral interarea anacline. Sulcus originating 
near mid-valve, strongly deepening anteriorly, flanked laterally by 
angular plications. Dorsal valve moderately convex with swollen 
umbo. Strong median fold with steep sides, originating anteriorly of 
mid-valve, bisected medianly by fine groove. Shell surface smooth 
with rare, slightly irregular growth lamellae anteriorly. Ventral inte- 
rior with teeth supported by short subparallel dental plates. Dorsal 
interior with forked cardinal process on a short shaft. 

Measurements. 
T=4.5. 



(488/12375) dorsal valve, L=16.2, W=16.9, 



Discussion. These shells are comparable with large specimens of 
Triplesia aff. subcarinata, but differ in having a narrow groove 
bisecting the dorsal median fold and a ventral sulcus which is more 
rounded in cross-section and a dorsal median fold. However, the 
shape of the dorsal fold and ventral sulcus varies significantly and it 
is difficult to evaluate observed morphological differences in this 
species because of the small number of shells available. 

Triplesia aff. subcarinata Cooper, 1956 

PI. 10, figs 1-8, 19 

Material. 6 conjoined valves, 2 ventral and 3 dorsal valves from 
Samples 100 (=K-98/1970), 626 (BC 57515), 8251a (BC 57514), 
Anderkenyn-Akchoku section; Samples 85258 (BC 57094-5), 
Kujandysai section; Sample 1041a, Burultas Valley. 

Discussion. This species resembles Triplesia carinata Cooper, 
1956, from the Pratt Ferry Formation of Alabama and T subcarinata 
Cooper, 1956, from the Lebanon Formation of Tennessee, as well as 
specimens of that species from the Bestamak Formation (Llandeilo- 
Lower Caradoc) of the Chingiz range. Kazakhstan (Nikitin & Popov 
in Klenina et at. 1984), in having a well-developed sulcus and a 
carinate dorsal median fold originating near the mid length, but it 
differs from both in having a larger shell, a strongly convex lateral 
profile in the dorsal valve with maximum height near the anterior 
margin, and a variable transverse profile of the dorsal median fold 
which has a tendency to be rounded in most specimens. Triplesia aff. 
subcarinata differs from T ainca Severgina, 1978, from the Lower 
Ashgill Gurianovka Formation and Marinikha Limestone of the 
Sayano-Altai Mountain Region in having a strongly dorsibiconvex 
lateral profile with a slightly accentuated ventral beak erect 
posteriorly, a narrow ventral sulcus and a dorsal median fold usually 
rounded in cross-section. 



Genus BICUSPINA Havh'cek, 1950 

TYPE SPECIES. Orthis cava Barrande, 1848, from the Lower 
Caradoc of Bohemia. 



54 



L.E. POPOV, L.R.M. COCKS AND I.F. NIKITIN 




UPPER ORDOVICIAN BRACHIOPODS FROM KAZAKHSTAN 



55 



Bicuspina rukavishnikovae Klenina, 1984 PI. 9, figs 5-13 

1984 Bicuspina rukavishnikovae Klenina in Klenina et al.\ 62, pi. 
5, figs 7, 8. 

1985 Bicuspina attrita Popov: 61 , pi. 3, figs 2-A. 

Holotype. IGNA 41 1/89, conjoined valves, from the Abai For- 
mation (Lower Caradoc), Ordotas Mountains, Chingiz Range, 
Kazakhstan. 

Material. Five conjoined valves, one ventral and 9 dorsal valves 
from Samples 100 (=K-98/1970) (BC 57505), 626, Anderkenyn- 
Akchoku section; Sample 1018 (including the holotype of B. attrita 
CNIGR 32/1 1989), area 7 km southwest of Karpkuduk well, Kotnak 
Mountains. 

Description. Shell dorsibiconvex, about half as thick as long and 
about 90% as long as wide. Hinge line short, about two-thirds 
maximum width. Ventral valve gently convex in lateral profile with 
maximum thickness at about one-third valve length. Beak small, 
slightly curved. Ventral interarea low, planar, apsacline with small 
pseudodeltidium bisected by monticulum. Ventral median sulcus 
originating in umbonal area with flattened bottom and steep lateral 
sides about 60% valve width. Well-developed semioval tongue. 
Dorsal valve strongly convex with maximum thickness at mid- 
length and flattened umbonal area. Median fold high, originating 
near umbo, with steep lateral slopes. Radial ornament costellate with 
6-1 1 ribs in the fold and sulcus and 1 3-2 1 ribs on the lateral sides of 
mature shells. 

Ventral interior with small teeth supported by short diverging 
dental plates. Ventral muscle field open anteriorly, weakly im- 
pressed. Umbonal area with a short internal pedicle tube (PI. 9, 
fig. 10). Dorsal interior with forked cardinal process on a short, 
thickened shaft, and small curved socket ridges. 

Measurements. (47 1/1 2375) conjoined valves, L=14.0,W=1 6.9, 
T=6.3, Sw=8.9; (474/12375) conjoined valves, L=20.4, W=21.0, 
T=11.2, Sw=8.7; (475/12375) conjoined valves, L=22.9, W=24.5, 
T=10.2, Sw=12.8; (476/12375) conjoined valves, L=16.5, W=19.8, 
T=6.5, Sw=11.2; (479/12375) conjoined valves, L=10.5, T=6.7, 
Sw=6.7. 

DISCUSSION. Coarsely ribbed triplesiides are characteristic of the 
Caradoc of West Gondwana (Havlicek 1950; Melou 1990) and 
Avalonia (Williams 1963; 1974), but they are apparently absent from 
China and Australia and very rare in Kazakhstan. Specimens from 
the Chu-Ili Range were previously known as Bicuspina attrita 
Popov (1985). They differ from Bicuspina rukavischnikovae, 
described by Klenina {in Klenina etal. 1984) from the Abai Forma- 
tion of the Chingiz Range, only in having a slightly uneven lateral 
profile of the ventral valve with maximum height posterior to mid- 
length, and in a more apsacline ventral interarea. These differences 



are regarded here as intraspecific, and specimens of Bicuspina from 
the Chu-Ili and Chingiz ranges are therefore conspecific. Klenina 
mentioned the presence of a forked cardinal process and internal 
pedicle tube in the original description of the species, but the 
interiors of both valves were not illustrated. 

The published Llanvirn age of B. rukavischnikovae in the Ordotas 
Mountains of the Chingiz Range, which is the type locality, is not 
supported by analysis of the associated brachiopod assemblage. It 
co-occurs with Hesperorthis karaadirensis Klenina, which is prob- 
ably synonymous with Paralenorthis numerosa (Nikitin & Popov) 
and rhynchonellids, suggesting that the age of the assemblage is not 
older than Llandeilo to early Caradoc. 



Genus GRAMMOPLECIA Wright & Jaanusson, 1993 

Type SPECIES. Grammoplecia triplesioides Wright & Jaanusson, 
1993, from the Boda Limestone (Ashgill) of Dalarna, Sweden. 



Grammoplecia wrighti sp. nov. 



PI. 9, figs 14-21 



ETYMOLOGY. After A. D. Wright, to honour his triplesioid studies. 

HOLOTYPE. BC 57509, PI. 9, figs 17, 18, a dorsal valve from the 
Anderken Formation, Sample 8214, Anderkenyn-Akchoku section. 

Material. Three pairs of conjoined valves, three ventral and 9 
dorsal valves, from Samples 620 (BC 57100-04, 57506, 07, 10, 1 1), 
626 (BC 57105-6), Anderkenyn-Akchoku section; Samples 8214 
(BC 57099, 57508-11), 8215b (BC 57107), west side of Ashchisu 
River; Sample 628, Kujandysai Section,. 

DESCRIPTION. Shell dorsibiconvex, slightly transverse, sub- 
rectangular to suboval in outline, about 83% as long as wide. Hinge 
line straight, not exceeding two-thirds shell width. Anterior commis- 
sure uniplicate. Ventral valve moderately convex with low, apsacline 
interarea and flat pseudodeltidium bisected by monticulum. Ventral 
sulcus originating about 3-5 mm from the umbo, ending in wide, 
trapezoidal tongue about 83% as wide as the valve. Dorsal valve 
strongly convex with maximum thickness at about one-third valve 
length. Strong dorsal median fold, flat centrally with steep lateral 
slopes. Lateral slopes convex in cross-section, strongly inclined to 
the commissural plane. Radial ornament of fine capillae about 5-7 
per mm crossed by fine, closely-spaced concentric fila. 

Ventral interior with strong teeth and long, diverging, widely 
spaced dental plates. Ventral muscle field large, about two-fifths 
valve length, slightly raised anteriorly with wide, subtriangular 
adductor track dividing narrow, strip-like diductor scars. Dorsal 
interior with forked cardinal process on strong short shaft joined to 
low and short socket plates. Adductor field quadripartite with ante- 
rior and posterior pairs separated by strong, transverse ridges. Low 



PLATE 9 

Figs \-<\ Gacella institata sp. nov.. Sample 100, Anderkenyn-Akchoku section. 1, BC 57498, ventral valve, umbonal area, x 5. 2, Sample 100, BC 57502. 

enlargement of interarea of conjoined valves, x 3. 3, 4, BC 57499, dorsal valve exterior, x 2, and umbonal area showing radial ornament, x 6. 
Figs 5-13 Bicuspina rukavishnikovae Klenina, 1984. 5-9. Sample 100, Anderkenyn-Akchoku section, BC 57505, conjoined valves, dorsal, lateral, ventral 

anterior and posterior views, x 2. 10-13, Sample 1018, area 7 km southwest of Karpkuduk well, Kotnak Mountains; 10, CNIGR 31/1 1989. ventral 

internal mould showing internal pedicle tube, x 5; 11, CNIGR 32/1 1989, dorsal exterior, x 2: 12, 13, CNIGR 33/1 1989, conjoined valves, ventral view, x 

2, posterior view, x 5. 
Figs 14-21 Grammoplecia wrighti sp. nov. Anderkenyn-Akchoku section. 14, 15, 19-21, Sample 620; 14, BC 57506. ventral internal mould, x 2; 15. BC 

57507, conjoined valves, oblique posterior view of internal mould showing cardinal process, x 2; 19,20, BC 57510, dorsal and lateral views of exterior, x 

2; 21, BC 5751 1 , dorsal internal mould, x 2. 16-18, Sample 8214; 16, BC 57508, dorsal exterior, x 2; 17, 18. BC 57509. holotype, dorsal exterior and 

lateral views, x 2 
Figs 22-26 Triplesia sp. Sample 8228, east side of Kopalysai. 22, 23, 25. dorsal internal mould, lateral and dorsal views, x 2. posterior view showing 

cardinal process, x 3. 24, BC 57512. latex cast of dorsal exterior, x 2. 26. ventral internal mould, x 2. 



56 



L.E. POPOV, L.R.M. COCKS AND I.F. NIKITIN 



0.45 0.60 0.75 \ 





1.35 



1.50 



5 mm 



1.80 



Fig. 13 Transverse serial sections of Placotriplesia spissa sp.nov., BC 57605 from Sample 628. Distance in mm is measured from the posterior tip 
of ventral beak. Dorsal valve uppermost. 



and narrow median ridge extending to anterior border of the muscle 
field. 

Measurements. (490/12375) dorsal valve, L=17.2, W=19.2, 
T=8.2, Sw=9.6; (491/12375) dorsal valve, L=17.2, W=21.0, T=8.6. 
Sw=8.2; (495/12375) dorsal valve, L=10.2,W=15.3,T=7.8,Sw=4.6. 

Discussion. This species differs from Grammoplecia triple sioides 
Wright & Jaanusson, 1993, G. globosa (Nikitin & Popov, 1985), 
from the Andryushinka Formation (Llandeilo-Lower Caradoc) of 
north-central Kazakhstan, G. krotovi (Chernyschev, 1887) from the 
Upper Caradoc to Lower Ashgill of Novaya Zemlya, Vaigach and the 
Urals (Bondarev 1968) and G. sibirica (Nikiforova, 1955) from the 
Upper Caradoc of Siberia, in having a more transverse shell outline, 
a ventral sulcus and dorsal median fold with steep lateral sides and 
flat centrally, a strongly and evenly convex dorsal profile with 
maximum height at mid-length and a broader hinge line. 



Genus PLACOTRIPLESIA Amsden, 1968 

Type SPECIES. Triplesia praecipta Ulrich & Cooper, 1936a, from 
the Wenlock of Arkansas, U.S.A. 

Placotriplesia spissa sp. nov. 



Holotype. BC 57517, PI. 10, figs 9-13, from Sample 2538, 
Anderken Formation, Akchoku Mountain, Kujandysai section. 

Material. 9 pairs of conjoined valves. 6 ventral and 12 dorsal 
valves from Samples 8214, 8215, 8223, Anderkenyn- Akchoku sec- 
tion; Samples 628, 2538 (BC 575 1 7 ), 82 1 9, 8256, Kujandysai section. 

Description. Shell smooth, dorsibiconvex profile, about 80% as 
thick as long and 75% as long as wide, transverse and suboval in 
outline, with maximum width at mid-length. Hinge-line short, less 
than half valve width. Anterior commissure strongly uniplicate. 
Ventral valve moderately convex, with an erect beak and minute 
apical foramen. Ventral interarea high, planar, apsacline with flat 
pseudodeltidium. Ventral sulcus originating from quarter to half 
valve length, strongly deepening anteriorly, with strong geniculated 
tongue about 75% valve width, and inclined at less than a right angle 
towards commisural plane. Lateral sides of sulcus accentuated by 
angular plications. Dorsal valve strongly convex, with swollen 
incurved beak; dorsal median fold strong and rounded in cross- 
section. Ventral interior with delicate teeth and short divergent dental 
plates. Dorsal interior with grooved forked cardinal process with 
strongly curved prongs posteriorly with distal parts subparallel to 
commisural plane, separated proximally and fused with narrow 
curved socket ridges. 

PI. 10, figs 9-18; Figs 13, 15 Discussion. This species represents the earliest record of 



PLATE 10 

Figs 1-8, 19 Triplesia aff. subcarinata Cooper. Anderkenyn- Akchoku section. 1-4, 19, Sample 825 la, BC 575 14, conjoined valves, ventral, dorsal, posterior 
and anterior views, x 2, dorsal view of the umbonal area showing pseudodeltidium with monticulus, x 4. 5-8, Sample 100, BC 575 15. conjoined valves, x 2. 

Figs 9-18 Placotriplesia spissa sp.nov. 9-14, Sample 2538, Akchoku Mountain, Kujandysai section. 9-13, BC 575 1 7. holotype, conjoined valves, 
ventral, dorsal, anterior, posterior and lateral views, x 2. 14, CNIGR 3/12361, conjoined valves showing ventral interarea with pseudodeltidium lacking 
monticulum, x 5. 15-18, Sample 8214. Anderkenyn-Akchoku section, CNIGR 3/12361. conjoined valves, ventral, dorsal, anterior and lateral views, x 2. 

Figs 20-22 Skenidioides sp. Sample 8214, west side of Ashchisu River, BC 57518, conjoined valves, ventral, dorsal and posterior views, x 6. 

Figs 23-29, 31-36 Dolerorthis pristina sp. nov. 23-25, 32, 34. Sample 626. Anderkenyn-Akchoku section. 23-25, 32. BC 57519, conjoined valves, 
ventral, lateral and dorsal views, x 2, detail of the shell surface, x 8; 34, BC 57522, ventral internal mould, x 2. 26-29, 31, Sample 620, Anderkenyn- 
Akchoku section, BC 57520. holotype, conjoined valves, dorsal, ventral, lateral, posterior and anterior views, x 2. 33, 35, Sample 2538, Akchoku 
Mountain, Kujandysai section; 33. BC 57521. ventral exterior, x 2; 35, BC 56768, dorsal interior, x 3. 36, Sample 8214. Anderkenyn-Akchoku section, 
BC 57523, dorsal internal mould, x 2. 

Figs 37^12 Glyptorthis sp., Kujandysai section. 37, Sample 2538, BC 57524, ventral internal mould, x 3. 38^12, Sample 628 (=K- 107/70), west side of 
Kujandysai, BC 57525, conjoined valves, anterior, dorsal, lateral and ventral views, x 3, dorsal view, x 6. 

Fig. 43 Plectorthis? bundtasica sp. nov. Sample 1018, area 7 km SW of Kotnak mountains, south Betpak-Dala, CNIGR 2/1 1989, latex cast of ventral 
interior, x 2. 



UPPER ORDOVICIAN BRACHIOPODS FROM KAZAKHSTAN 



57 




58 



L.E. POPOV, L.R.M. COCKS AND I.F. NIKITIN 



Table 19 Measurements of complete shells of Placotriplesia spissa sp. nov.. Sample 2538. Kujandysai section, 822 1 from Anderkenyn-Akchoku section 
and Sample F- 104 la from Burultas valley. 





L 


W 


T 


Sw 


St 


LAV 


T/L 


SwAV 


St/Sw 


N 


7 


7 


7 


7 


3 


7 


7 


7 


3 


X 


13.9 


19.0 


11.5 


10.4 


7.7 


73.9% 


81.6% 


75.5% 


73.2% 


s 


2.61 


4.27 


3.13 


1.43 


3.04 


7.9 


12.7 


6.1 


22.3 


MIN 


10.5 


13.6 


6.8 


8.2 


3.3 


60.4% 


61.7% 


67.9% 


37.5% 


MAX 


16.3 


24.5 


14.7 


11.5 


7.6 


83.9% 


90.2% 


83.0% 


69.1% 



Placotriplesia, which is otherwise known only from the Silurian. It 
differs from P. praecipta, P. juvenis Ulrich & Cooper (1936a), P. 
waldronensis (Miller & Dyer), and P. rostellata (Ulrich & Cooper 
1936a) in having a strongly dorsibiconvex shell, high dorsal median 
fold and deep ventral median sulcus originating at some distance 
from the umbo. It also differs from P. praecipta, as redefined by 
Amsden (1968). in having a strongly curved cardinal process with 
posteriorly-directed prongs. 



Order PROTORTHIDA Schuchert & Cooper, 1931 

Superfamily PROTORTHOIDEA Schuchert & Cooper, 1931 

Family SKENIDHDAE Kozlowski, 1929 

Genus SKENIDIOIDES Schuchert & Cooper, 1931 

TYPE SPECIES. Skenidioides billingsi Schuchert & Cooper, 1931, 
from the Caradoc of Ontario, Canada. 



Skenidioides sp. 



PI. 10, figs 20-22 



Material. One pair of conjoined valves (L=2.7. W=5.4, T=1.7), 
one ventral and one dorsal valves from Samples 8223b and 8226, 
Anderkenyn-Akchoku section; Sample 8214 (BC 57518), west side 
of Ashchisu River. 

Discussion. The shells from the Anderken Formation closely 
resemble Skenidioides anthonensis (Sardeson). as redescribed and 
illustrated by Cooper (1956: 491 ), in the general shape of the shell, 
narrow median sulcus in the dorsal valve, carinate ventral valve and 
characters of radial ornament, but differ somewhat in having a more 
flattened dorsal valve. Although the exterior is characteristic of 
Skenidioides. the absence of known interiors in the specimens from 
the Anderken Formation precludes specific identification. 



Order ORTHIDA Schuchert & Cooper, 1932 

Suborder ORTHIDINA Schuchert & Cooper, 1932 

Superfamily ORTHOIDEA Woodward, 1852 

Family HESPERORTHIDAE Schuchert & Cooper, 1931 

Genus DOLERORTHIS Schuchert & Cooper, 1931 

Type species. Orthis interplicata Foerste, from the Niagara Group 
(Silurian) of Indiana, U.S.A. 

Dolerorthis expressa Popov, 1980 

PI. 1, fig. 29, PI. 11, figs 1,2 

1980 Dolerorthis expressa Popov: 144, pi. 1, figs 5-7. 

Holotype. CNIGR 11/11 523, ventral internal and external moulds 
(L=18.4, W=24.7), from the Anderken Formation, Sample 1018. 7 
km southwest of Karpkuduk well, Kotnak Mountains. 

MATERIAL. One pair of conjoined valves, 1 2 ventral and 1 1 dorsal 
valves, internal and external moulds, from Sample 8 1 37 (BC 57526), 



Anderkenyn-Akchoku; Sample 817, about 4 km south-west of Alakul 
Lake; Sample 1018 (BC 57368),7 km southwest of Karpkuduk well, 
Kotnak Mountains, south Betpak-Dala. 

Description. Shell subequally biconvex, transverse, subrect- 
angular in outline, about 15% as long as wide with maximum width 
anterior to hinge line. Cardinal extremities slightly rounded; anterior 
commissure weakly unisulcate; ventral valve gently convex in lat- 
eral profile with maximum thickness at about one-third from anterior 
margin; ventral interarea apsacline, slightly curved in cross-section 
with open triangular delthyrium. Dorsal valve moderately convex 
with shallow sulcus originating at the umbo. Interarea low, planar, 
anacline. Radial ornament variably multicostellate with costellae of 
two to three generations. 4-6 ribs per 3 mm along the posterior 
margin of adult specimens. Concentric ornament of fine, ridge-like, 
evenly spaced fila, 3^4 per mm. 

Ventral interior with strong teeth supported by diverging dental 
plates continuing anteriorly as elevated muscle bounding ridges 
enclosing an elongate, subrhomboidal muscle field about two-fifths 
as long as the valve. Adductor scars narrow, strip-like, completely 
separating large, deeply impressed diductor scars of about equal 
length. Mantle canals saccate with subparallel to slightly converging 
vascula media. Dorsal interior with simple, ridge-like cardinal proc- 
ess on the high notothyrial platform slightly inclined posteriorly. 
Brachiophores high, triangular with slightly diverging bases. Weakly 
impressed dorsal adductor scars divided posteriorly by a very short 
median ridge. 

Discussion. This species is somewhat similar to Dolerorthis 
tenuicostata Williams (in Whittington & Williams 1955: 406) from 
the Lower Caradoc of Wales, but differs in having a more trans- 
verse shell outline, lateral profile of the ventral valve with 
maximum thickness anterior to the mid-length in full grown speci- 
mens and a weak dorsal sulcus continuing towards the anterior 
margin. It differs from Dolerorthis aff. hubeiensis Zeng. which 
occurs in the Dulankara Regional Stage of north Betpak-Dala, 
Kazakhstan (Nikitin et al. 1996), in having a subequally biconvex 
transverse shell, finer radial ornament and more widely spaced 
concentric fila. 

Dolerorthis pristina sp. nov. PI. 10, figs 23-29, 31-36 

ETYMOLOGY. After pristinus, Latin - former. 

Holotype. BC 57520, PI. 10, figs 26-29, 31, conjoined valves 
(L=9.7. W= 1 2.6, T=4.2) from the Anderken Formation, Sample 620, 
Anderkenyn-Akchoku section. 

Material. 8 pairs of conjoined valves, 15 ventral and 18 dorsal 
valves from Samples 100 (=K-98/1970) (BC 57110-7), 620 (BC 
57520), 626 (BC 57130-32, 57519), 8223a (BC 57158, 59), 8223b, 
Anderkenyn-Akchoku; Sample 8214 (BC 57150-54, 57523), 
Ashchsu River; Samples 628 (BC 57133-5), 2538 (BC 56768, 
57141^15, 57521, 22), 8217 (BC 57156-7), Kujandysai near 
Akchoku Mountain; Sample 948 (BC 57136-40), Tesik River. 



UPPER ORDOVICIAN BRACHIOPODS FROM KAZAKHSTAN 

Table 20 Measurements of complete shells of Dolerorthis pristina sp. 
nov.. Samples 100 and 626 from Anderkenyn-Akchoku section. 





Lv 


W 


T 


Iw 


Lv/W 


T/L 


IwAV 


N 


10 


10 


6 


10 


10 


6 


9 


X 


10.0 


11.8 


4.2 


10.3 


85.2% 


47.6% 


87.9% 


s 


3.04 


3.36 


1.11 


2.63 


10.0 


6.5 


9.5 


MIN 


5.9 


7.5 


2.8 


6.5 


70.2% 


40.0% 


75.0% 


MAX 


14.2 


18.2 


5.5 


13.7 


103.6% 


56.3% 


100.0% 



Description. Shell weakly ventribiconvex, slightly transverse, 
suboval in outline, on average 82% (S=7.0, N=10) as long as wide 
and 48% (S=6.0, N=6) as high as long. Anterior commissure slightly 
shorter than maximum shell width at mid-length. Cardinal extremi- 
ties obtuse. Anterior commissure rectimarginate or weakly sulcate. 
Radial ornament costellate with up to 1 7 primary costae and costellae 
branching near the mid-length and near the anterior and lateral 
margins. 6-8 costellae per 3 mm along the anterior margin and 
varying from 31 to 54 in full-grown specimens. 

Ventral valve moderately and unevenly convex with maximum 
thickness at about one-third valve length from the small, pointed 
beak. Ventral interarea apsacline, slightly curved in cross-section 
with open, narrow delthyrium. Dorsal valve weakly convex with 
maximum thickness slightly anterior from the beak. Interarea low, 
planar, linear. Shallow sulcus usually well defined in the posterior 
half of the dorsal valve, but fading anteriorly. 

Ventral interior with small teeth and low, divergent dental plates. 
Muscle field small, slightly elongate, subpentagonal in outline. 
Adductor scars narrow, completely separating diductor scars of 
about equal length. Mantle canals saccate with slightly divergent 
proximal parts of vascula media. Dorsal valve interior with high, 
subtriangular brachiophores diverging anteriorly. Cardinal process 
ridge-like with crenulated myophore. situated on a low subtriangular 
notothyrial platform. Adductor muscle field subrectangular with 
anterior adductor scars slightly larger than posterior. Median ridge 
low and broad, bisecting the entire adductor muscle field. 

DISCUSSION. Dolerorthis pristina differs from D. expressa (Popov 
1980) in its less convex dorsal valve, much smaller size (less than 
half D. expressa), and in its evenly convex ventral profile, in contrast 
to D. expressa in which the ventral profile is relatively flat near the 
umbo, but increases greatly anteriorly. In addition, D. pristina has 
finer radial ornament. 

Zeng (1987) erected Paradolerorthis as a subgenus within 
Dolerorthis. However, his quoted distinctions and equivocal illustra- 
tions do not allow us to recognize his subgenus as useful, but the type 
species D. (Paradolerorthis) calla appears similar to D. pristina. 



Family GLYPTORTHIDAE Schuchert & Cooper, 
Genus GLYPTORTHIS Foerste, 1914 



1931 



Type species. Orthis insculpta Hall, 1 847, from the Richmondian 
(Ashgill), New York, U.S.A. 



Glyptorthis sp. 



PI. 10, figs 37-42 



Material. Five pairs of conjoined valves, 7 ventral and 6 dorsal 
valves from Samples 620 (BC 57163, 64), 8223a, Anderkenyn- 
Akchoku; Sample 8214, Ashchisu River; Samples 2538 (BC 
57 166-69, 57524), 8256, Kujandysai near Akchoku Mountain; Sam- 
ple 628 (BC 57165, 57525), east side of Kujandysai; Sample 948, 
Tesik River. 

Description. Shell slightly ventribiconvex, transverse, rounded 



59 

subrectangular in outline, about 80% as long as wide. Hinge line 
slightly narrower than maximum shell width at mid-length. Cardinal 
extremities slightly obtuse. Anterior commissure varying from 
slightly sulcate to rectimarginate. Ventral valve moderately convex 
with maximum thickness at umbo. Interarea moderately high, trian- 
gular, planar, catacline, divided by narrow triangular, open delthyrium. 
Dorsal valve gently convex with maximum thickness at about one- 
quarter valve length from the beak. Dorsal sulcus shallow and 
narrow, originating at umbo and fading anteriorly. Dorsal interarea 
low, linear, orthocline. Radial ornament coarsely costellate with up 
to 16 primary ribs and 25-30 costellae (up to 5 costellae per 3 mm) 
in adult specimens. Secondary costellae in the median part of the 
dorsal valve bifurcate internally. Concentric ornament of crowded, 
evenly spaced growth lamellae. 

Ventral interior with teeth supported by short and high dental 
plates. Muscle field small, situated entirely within delthyrial cham- 
ber. Mantle canals saccate with straight, slightly diverging anteriorly 
vascula media. Dorsal interior not observed. 

DISCUSSION. These specimens closely resemble Glyptorthis 
balcletchiensis (Davidson, 1883) from the Upper Caradoc of the 
Girvan District, Scotland (Williams 1962) in the size, general outline 
and convexity of the shell, as well as in the number and bifurcation 
of the costellae. It differs from another coarsely ribbed Kazakh 
species Glyptorthis? bestamaki Nikitin & Popov (in Klenina et al. 
1984) from the lower Bestamak Formation (Nemagraptus gracilis 
Zone) of the Chingiz Range in having a rectimarginate or slightly 
sulcate anterior commissure and a dorsal sulcus not reversed anteriorly 
into the median fold. 



Family PLAESIOMYIDAE Schuchert, 1913 
Genus AUSTINELLA Foerste, 1909 

Type SPECIES. Orthis kankakensis McChesney, from the 
Maquoketa Formation (Ashgill) of Iowa, U.S.A. 

Austinella sarybulakensis sp. nov. PI. 11, figs 15-22 

Etymology. After Sarybulak River, 10 km west of the type locality. 

HOLOTYPE. BC 56507, PI. 11. figs 15-18, conjoined valves; 
Anderken Formation, Sample 85258, east side of Uzunbulak River. 

Material. Three conjoined valves, one ventral and one dorsal 
valve from Sample 85258 (BC 56505-8), east side of Uzunbulak 
River; Sample 818a, Burultas Valley. 

Description. Shell subequally biconvex, transverse, subrect- 
angular in outline about 93-97% as long as wide and 54-60% as 
thick as long. Hinge line somewhat shorter than maximum shell 
width at the mid-length. Anterior commissure uniplicate. Ornament 
costellate with 8-9 costellae per 5 mm along the anterior margin and 
25-28 primary ribs near the umbo. Ventral valve moderately convex 
with maximum thickness slightly anterior to the umbo. Ventral 
pseudointerarea high, triangular with open, triangular delthyrium. 
Shallow sulcus originating at about 7 to 9 mm anterior to the beak, 
widening and deepening anteriorly. Lateral sides of the valve gently 
and evenly convex in transverse section. Dorsal valve moderately 
and unevenly convex. Dorsal interarea low, planar, orthocline. Shal- 
low dorsal sulcus in the umbonal area reverses into a median fold at 
5-7 mm from the umbo. 

Ventral interior with strong teeth and short, slightly divergent 
dental plates. Muscle field strongly raised anteriorly in a form of 
pseudospondylium, rounded subtriangular in outline, about two- 



L.E. POPOV, L.R.M. COCKS AND I.F. NIKITIN 




UPPER ORDOVICIAN BRACHIOPODS FROM KAZAKHSTAN 



61 



fifths as long as the valve. Mantle canal system saccate with short, 
diverging vascula media. Dorsal interior with blade-like cardinal 
process situated on a high subtriangular notothyrial platform. 
Brachiophores high, subtriangular with strongly thickened, slightly 
divergent bases. Adductor field weakly impressed, quadripartite, 
bisected by a low median ridge. Anterior adductors larger than 
posterior ones. 

Measurements. Conjoined valves; [BC 56507, L=22.2, W=24.0. 
T=12.2, Sw=14.4, St=6.4; BC 56508, L=25.1, W=26.3, T=14.8. 
Sw=14.6, St=6.8]. 

DISCUSSION. This species can be distinguished from other species 
of the genus such as Austinella whitfieldi (Winchell & Schuchert) 
and A. kankakensis (McChesney), re-described by Wang (1949) 
from the Ashgill Maquoketa Formation of Iowa, by its uniplicate 
anterior commissure with the dorsal sulcus reversed into the median 
fold posterior to the mid-valve. 



Superfamily PLECTORTHOIDEA Schuchert, 1929 

Family PLECTORTHIDAE Schuchert, 1929 

Subfamily PLECTORTHINAE Schuchert, 1929 

Genus PLECTORTHIS Hall & Clarke, 1892 

TYPE SPECIES. Orthis plicatella Hall, from the Cincinnatian (Lower 
Ashgill), of the U.S.A. 

Plectorthisl burultasica sp. nov. PL 10, fig. 43, PL 11, figs 
3-12 

1985 Plectorthis afi. a/to'ra Severgina; Popov: 51, pi. 1, figs 1,2. 

ETYMOLOGY. After the type locality in the Burultas Valley. 

HOLOTYPE. BC 57528, PL 11, figs 3^1, dorsal valve, Anderken 
Formation, Sample 7613, Akchoku Mountain, Kujundysai section. 

Material. Five conjoined valves, 8 ventral and 9 dorsal valves 
from Samples 2538, 7613 (BC 57528), Kujandysai near Akchoku 
Mountain; Sample 626 (BC 57170, 71), 843, 8128 (BC 57530), 
Anderkenyn-Akchoku; all Chu-Ui Range; Sample 1041a (BC 57529), 
Burultas Valley; Sample 1018 (BC 57367), 7 km southwest of 
Karpkuduk well, Kotnak mountains, south Betpak-Dala. 

DESCRIPTION. Shell subequally biconvex, transversely subrect- 
angular in outline, about 60% as thick as long and three-quarters as 



long as wide, with maximum width at mid-length. Cardinal extremi- 
ties obtuse to slightly rounded. Anterior commissure rectimarginate. 
Ventral valve moderately and gently convex in lateral profile with 
low, strongly apsacline interarea slightly curved in cross-section. 
Dorsal valve moderately and evenly convex in lateral profile with 
low, linear, orthocline interarea and shallow umbonal sulcus fading 
at mid-length. Radial ornament of 28-32 rounded costellae bifurcat- 
ing at the posterior half of the shell and separated by interspaces of 
about equal width as ribs. Radial rows of fine rounded exopunctae on 
both sides of each rib. 

Ventral valve with teeth supported by thin diverging dental plates 
about one-third shorter than length of the elongate subpentagonal 
muscle field. Adductor scar narrow triangular, slightly raised 
anteriorly, about the same length as strongly impressed diductor 
scars. Dorsal valve interior with high, triangular brachiophores with 
bases converging anteriorly. Notothyrial platform high and narrow, 
crossed by ridge-like cardinal process with crenulated myophore. 
Median ridge strong, extending anteriorly to mid-valve. Dorsal 
anterior and posterior adductor scars about equal size, separated by 
fine, slightly oblique, transverse ridges. 

Discussion. Criteria for generic separation amongst the 
Plectorthidae require revision. The type species of Plectorthis, P. 
plicatella, has never been fully revised, although Schuchert & Cooper 
(1932, pi. 11, figs 4, 9) illustrated ventral and dorsal interiors from 
the Maysville Formation of Cincinnati, Ohio. Material labelled as P. 
plicatella in the Natural History Museum (eg. BB 14835) from 
Cincinnati includes well preserved exteriors with no trace of 
exopunctae. However, our new species has well-developed rows of 
exopunctae on both sides of each costa, as have other species 
attributed to Plectorthis, e.g. Plectorthisl punctata illustrated by 
Cooper (1956), P. obesa, mentioned but not illustrated by Cooper 
(1956), and the Plectorthis sp. of Neuman (1971:21). Thus, until 
comparable exopunctae have been found in true P. plicatella, we 
attribute the exopunctate species to Plectorthis with a query. 

Plectorthisl burultasica resembles Plectorthis altaica Severgina, 
1967 from the Khankhara Formation (Caradoc) of Gornyi Altai in 
radial ornament, size and outline, but differs in having a weak 
umbonal dorsal sulcus fading to the mid-valve and a more 
ventribiconvex profile. Plectorthisl punctata has a slightly sulcate 
dorsal valve by comparison with PI burultasica as well as fewer rib 
bifurcations. It is also considerably smaller. In addition, P.l obesa 
has a more strongly convex dorsal valve and fewer bifurcations in the 
ribbing. 



PLATE 11 

Figs 1, 2 Dolerorthis expressa Popov, Sample 8137, Anderkenyn-Akchoku section. BC 57526, dorsal internal mould, x 2. and latex of external mould, x 
6. 

Figs 3-12 Plectorthis? burultasica sp. nov. 3, 4. Sample 7613, Akchoku Mountain. Kujandysai section. BC 57528. holotype. dorsal latex cast and internal 
mould, x 2. 5-8, Sample 1041a, Burultas, BC 57529, conjoined valves, lateral, posterior, ventral and dorsal views, x 2. 9-11. Sample 1018, area 7 km 
SW of Kotnak mountains, south Betpak-Dala, CNIGR 1/1 1989, conjoined valves, dorsal, ventral and lateral views, x 2. 12, Sample 8128. Anderkenyn- 
Akchoku section, BC 57530. ventral exterior, latex cast, x 2. 

Figs 13, 14 Phaceloorthisl sp. Sample 2538, Akchoku Mountain, Kujandysai section, BC 57531, conjoined valves, ventral and dorsal views, x 1 .5. 

Figs 15-22 Austinella sarybulakensis sp. nov. Sample 85258, east side of Kujandysai. 15-18, BC 56507, conjoined valves, holotype, ventral, lateral, 
anterior and dorsal views, x 1.5. 19, BC 56508. conjoined valves, posterior view, x 1.5. 20. BC 56505, ventral interior, x 2. 21, 22, BC 56506, dorsal 
internal mould and latex cast, x 2. 

Fig. 23-37 Bowanorthisl devexa sp. nov. Sample 2538, Akchoku Mountain, Kujandysai section. 23-27, BC 57532, conjoined valves, holotype. ventral, 
lateral, anterior, dorsal and posterior views, x 3. 28-32, BC 57533, conjoined valves, lateral, dorsal anterior, posterior, and ventral views, x 3. 33-37, BC 
57534, conjoined valves, lateral, posterior, ventral, dorsal and anterior views, x 3. 

Figs 38-44 Phragmorthis conciliata Popov. 38. Sample 1018, area 7 km SW of Kotnak mountains, south Betpak-Dala. CNIGR 1 1/1 1989, ventral internal 
mould, x 4. 39, Sample 76 1 3, Akchoku Mountain. Kujandysai section. BC 57535, ventral internal mould, x 5. 40^12, Sample 2538, Akchoku Mountain, 
Kujandysai section, BC 57536, conjoined valves, lateral, dorsal and ventral views, x 4. 43, 44. Sample 626. Anderkenyn-Akchoku section. BC 57537. 
ventral exterior and lateral view, x 4. 



62 



L.E. POPOV, L.R.M. COCKS AND I.F. NIKITIN 



Genus PHACELOORTHIS Percival, 1991 

Type species. P. decoris Percival, 1991, from the Quondong 
Limestone (Caradoc) of New South Wales, Australia. 



Phaceloorthis? sp. 



PI. 11, figs 13, 14 



Material. One pair of conjoined valves (BC 57531) (L=20.8, 
W=25.6, T=9.8) from Sample 2538, Kujandysai, near Akchoku 
Mountain. 

Description. Shell subequally biconvex, transversely subrect- 
angular in outline with hinge line somewhat shorter than maximum 
shell width at mid-length. Anterior margin rectimarginate. Ventral 
valve moderately convex in lateral profile with maximum thickness 
at one-third valve length. Ventral interarea almost orthocline, slightly 
curved in cross-section with open, narrow triangular delthyrium. 
Dorsal valve moderately convex with maximum thickness near the 
mid-valve. Umbonal area with shallow sulcus fading towards mid- 
valve. Dorsal interarea anacline, planar, linear. Shell surface with 9 
low, angular radial plications and sumperimposed finely fascico- 
stellate ribs 7-9 per 3 mm along the anterior margin. Interior of both 
valves unknown. 

Discussion. The interior of this species remains unknown, which 
makes its generic attribution highly provisional, but there are only a 
few impunctate mid and late Ordovician orthide genera with 
fascicostellate ornament and they are mostly related to the family 
Giraldiellidae. Otherwise fascicostellate ribbing is characteristic of 
the plectorthid genus Phaceloorthis. This single specimen somewhat 
resembles Phaceloorthis recondita Popov, Nikitin & Cocks, 2000 
from the Otar Member (Upper Caradoc ) of the Dulankara Moutains in 
its general shell shape and size and fascicostellate ornament, but differs 
in having an orthocline ventral interarea and weak radial plications. 

Family WANGYUHDAE Zeng, 1989 
Genus BOWANORTHIS Percival, 1991 



Type species. Bowanorthis fragilis Percival, 1991, 
Caradoc of New South Wales, Australia. 



from the 



Bowanorthis! devexa sp. nov. PI. 11, figs 23-37; Fig. 14 

Etymology . After devexus, Latin - hollow. 

Holotype. BC 57532, PL 1 1 , figs 23-27, conjoined valves, from 
the Anderken Formation, Sample 2538, Kujandysai, near Akchoku 
Mountain. 

Material. Twelve conjoined valves from Samples 2538 (BC 
57178, 57532-34), 8217 (BC 57179) Kujandysai, near Akchoku 
Mountain; Sample 948 (BC 57172-77), Tesik River. 

Description. Shell ventribiconvex, slightly transverse and 
semioval, about 85-98% as long as wide with hinge line slightly 
shorter than maximum width at one-quarter shell length. Anterior 
commissure sulcate. Ventral valve carinate posteriorly, strongly 
convex in lateral profile with maximum thickness at about one-third 
valve length. Beak small, erect. Ventral interarea apsacline, incurved 
in cross-section, with small subtriangular delthyrium. Ventral me- 
dian fold originating at the umbonal area with steep lateral slopes 
flanked by folds. Lateral sides of the valves less convex. Dorsal valve 
evenly convex with linear, anacline interarea and deep, V-shaped 
median sulcus originating at the beak and ending in a low semioval 
tongue occupying slightly less than half of the valve length. Radial 
ornament fascicostellate with three strong angular ribs and 4-5 
costellae per mm along the anterior margin of mature specimens. 
Interior of both valves unknown. 

Measurements. Conjoined valves; (58/12375) L=6.4, W=6.7, 
T=3.8. Sw=3.3; (59/12375), L=4.5, W=4.6, T=2.7, Sw=2.6; (60/ 
12375), L=4.8,W=5.6,T=3.1,Sw=2.4. 

Discussion. This species is provisionally included within 
Bowanorthis because of the small ventribiconvex shell with strongly 
sulcate anterior commissure, carinate ventral valve, fascicostellate 
ornament and receding dental plates, all resembling B. fragilis from 
the Caradoc of New South Wales (Percival 1991 ); however, it differs 
in having a strongly apsacline ventral interarea and a less transverse 
shell outline. The general morphology of the cardinal process and 
brachiophores in the Kazakh species looks similar to B. fragilis, but 
the presence of the characteristic sigmoidal plates is impossible to 




Fig. 14 Transverse serial sections of Bowanorthis! devexa sp.nov. from Sample 2538. Kujandysai section. Distance in mm is measured from the posterior 
tip of ventral beak. Dorsal valve uppermost. Also reconstructions of the ventral and dorsal interiors. 



UPPER ORDOVICIAN BRACHIOPODS FROM KAZAKHSTAN 



63 



confirm from our sections (Fig. 14). In general shell shape and 
fascicostellate radial ornament the Anderken material is similar to 
Ranorthis Opik (1939) from the Volkhov and Kunda Stages of 
Baltica. However, the generic attribution of deve.xa is tentative and it 
may represent an undescribed genus. 

Family RANORTHIDAE Havlfcek, 1949 
Genus EODALMANELLA Havlfcek, 1950 

TYPE SPECIES. Orthis socialis Barrande, 1879; from the Sarka 
Formation (Llanvirn) of Bohemia. 

Eodalmanella extera Popov, 1985 PI. 12, figs 10-12, 15 

1985 Eodalmanella extera Popov: 54, pi. 1, figs 3-8. 

HOLOTYPE. CNIGR 4/11989, ventral internal mould (L=4.1. 
W=6.1), from the Anderken Formation, Anderkenyn-Akchoku sec- 
tion, Sample 843. 

Material. Five pairs of conjoined valves, 42 ventral and 48 dorsal 
valves, internal and external moulds, from Samples 100b, 620 (BC 
57189), 843, Anderkenyn-Akchoku section; Sample 1018, 7 km 
southwest of Karpkuduk well, Kotnak Mountains. 

DESCRIPTION. Shell ventribiconvex, transverse, subrectangular in 
outline, about 85% as wide as long with maximum width at mid- 
length. Anterior commissure gently sulcate. Ventral profile 
moderately convex, about one-third as thick as long, with maxi- 
mum thickness at about one-quarter valve length. Dorsal valve 
gently convex in transverse profile with maximum thickness 
slightly anterior to the umbo and with shallow median sulcus 
originating in the umbonal area. Dorsal interarea linear, strongly 
anacline. Radial ornament multicostellate with 3-6 ribs per mm 
along the anterior margin of mature specimens. Concentric orna- 
ment of very fine ridge-like, evenly spaced fila, often branching 
anteriorly. 

Ventral interior with teeth supported by short, divergent dental 
plates and rounded subtriangular, slightly elongate muscle field 
about 37% as long as the valve. Ventral adductor scars narrow, 
subtriangular, slightly shorter than diductor scars. Cardinal process 
ridge-like with bilobed, crenulated miophore. Brachiophores high, 
triangular, with widely diverging bases. Fulcral plates well devel- 
oped. Dorsal adductor scars quadripartite, extending anteriorly to 
mid-valve, bordered laterally by low ridges. 

DISCUSSION. This species strongly resembles Scaphorthis? aulacis 
Percival (1979a) from the Caradoc Goonumbla Volcanics of New 
South Wales in the impunctate shell, radial ornament and internal 
morphology of both valves. The Kazakh species differs in having 
longer dental plates, a cardinal process with a long shaft crossing all 
the bottom of the notothyrial cavity and well defined lateral ridges 
bordering the dorsal adductor field. A detailed discussion and basic 
statistics of this species were provided by Popov (in Nikitin & Popov 
1985). 

Table 22 Measurements of dorsal valves of Pionodema opima sp. nov., Sample 8228 from Kopalysai Section, sample 8229 from Buldukbai-Akehoku and 
Sample 7613 from Kujandysai section. 



Family CREMNORTHIDAE Williams, 1963 
Genus PHRAGMORTHIS Cooper, 1956 

Type species. Phragmorthis buttsi Cooper, 1 956, from the Effna- 
Rich Valley Formations (Llandeilo-Lower Caradoc) of Virginia, 
U.S.A. 

Phragmorthis conciliata Popov, 1985 

PI. 1 1, figs 38^14; PI. 12, figs 1-9 

1985 Phragmorthis conciliata Popov: 52, pi. 1, figs 9-12. 

HOLOTYPE. CNIGR 10/11989 (PI. 12, figs 1, 2), dorsal valve 
internal mould (L=4.5, W=7.2), from the Anderken Formation, 7 km 
south-west of Karpkuduk well, Kotnak Mountains, Sample 1018. 

Material. Two pairs of conjoined valves, 6 ventral and 8 dorsal 
valves, internal and external moulds, from Samples 626 (BC 57537), 
8223a, Anderkenyn-Akchoku section; Samples 2538 (BC 57536), 
7613 (BC 57535), Kujandysai section; Sample 8230, Buldukbai- 
Akehoku section; Sample 1018,7 km southwest of Karpkuduk well 
Kotnak Mountains; Sample 1024b, south side of Karatal River, south 
of Sorbulak spring. 

Diagnosis. Shell ventribiconvex, transverse, subrectangular out- 
line about 77% as wide as long with maximum width at mid-length, 
anterior commissure gently unisulcate; ventral valve strongly con- 
vex with maximum thickness between the umbo and mid-valve; 
ventral interarea high triangular, apsacline with narrow open 
delthyrium; dorsal valve moderately and evenly convex with narrow 
and shallow sulcus originating at the umbo; radial ornament finely 
and equally multicostellate with 5 ribs per mm along the anterior 
margin of mature specimens. Ventral interior with elongate 
subtriangular muscle field on pseudospondylium 2 1-26% as long as 
the valve. Dorsal interior with simple, ridge-like cardinal process on 
the high notothyrial platform which is strongly raised anteriorly; 
high, blade-like median septum 88% valve length; large, deeply 
impressed adductor scars, radially arranged, about 66% valve length. 

DISCUSSION. This species differs from Phragmorthis buttsi Cooper 
( 1956: 510) in its transverse subrectangular outline and lesser con- 
vexity of both valves. It is on average about half the size of 
Phragmorthis crassa Cooper (1956: 511) and has finer radial orna- 
ment. Both P. buttsi and P. crassa have a subcarinate ventral valve, 
which is another difference from the Kazakh species. 

Table 21 Measurements of ventral valves of Pionodema opima sp. nov.. 
Sample 8228 from Kopalysai Section, sample 8230 from Buldukbai- 
Akehoku and Sample 7613 from Kujandysai section. 





Lv 


W 


Ml 


Mw 


Lv/W 


Ml/L 


Iw/W 


N 


9 


9 


5 


6 


9 


5 


5 


X 


7.6 


9.4 


3.2 


3.5 


81.2% 


38.5% 


90.5% 


S 


1.38 


2.16 


0.63 


1.26 


8.6 


4.9 


14.9 


MIN 


5.0 


7.3 


2.6 


2.3 


68.5% 


34.2% 


64.9% 


MAX 


9.8 


13.8 


4.1 


5.7 


93.2% 


46.6% 


103.6% 





Ld 


W 


BB1 


BBw 


Ml 


Mw 


Lv/W 


Ml/L 


BBw/W 


N 


7 


7 


5 


5 


3 


3 


4 


3 


5 


X 


7.6 


9.5 


1.6 


2.6 


3.4 


3.1 


82.8% 


41.2% 


60.4% 


S 


2.36 


2.86 


0.15 


0.34 


0.32 


0.10 


10.7 


2.9 


9.9 


MIN 


3.2 


4.0 


1.4 


2.3 


3.2 


3.0 


70.1% 


38.8% 


50.0% 


MAX 


10.5 


12.9 


1.7 


3.1 


3.8 


3.2 


96.1% 


44.4% 


73.9% 



64 



L.E. POPOV, L.R.M. COCKS AND I.F. NIKITIN 



Suborder DALMANELLIDINA Moore, 1952 

Superfamily ENTELETOIDEA Waagen, 1884 

Family DRABOVIIDAE Havlicek, 1950 

Subfamily DRABOVIINAE Havlicek, 1950 

Genus PIONODEMA Foerste, 1912 

TYPE SPECIES. Orthis subaequata Conrad, from the Caradoc of Mis- 
souri, U.S.A. 

Pionodema opima sp. nov. PI. 12, figs 13, 14, 16-27 

ETYMOLOGY. After opimus, Latin - fat. 

Holotype. BC 57545, PI 12, figs 17-1 8, internal mould of conjoined 
valves, from the Anderken Formation, Sample 7613, Kujandysai sec- 
tion. 

Material. Nine pairs of conjoined valves, 70 ventral and 64 dorsal 
valves from Sample 7613 (BC 57545), Kujandysai section; 
Sample 8128, Anderkenyn-Akchoku; Sample 8228 (BC 57185, 57547, 
48), east side of Kopalysai; Samples 1 10, 8229 (BC 57546), 8230 (BC 
57803), 8257 (BC 57544), Buldukbai-Akchoku section; Sample 818a 
(BC 57802), Burultas Valley. 

Description. Shell slightly dorsibiconvex, transverse, suboval in 
outline, about 84% as long as wide and 53% as thick as long. Hinge line 
slightly shorter than the maximum shell width at mid-length. Anterior 
commissure uniplicate. Ventral valve gently convex with maximum 
thickness at the umbonal area. Beak curved, pointed and slightly erect 
posterior to hinge line. Ventral interarea subtriangular, apsacline, weakly 
curved in cross-section, with open triangular delthyrium. Shallow 
sulcus originating near mid-valve. Dorsal valve moderately convex 
with maximum thickness at quarter valve length. Umbonal area with 
shallow V-shaped sulcus reversed into a low and narrow median fold 
flanked laterally by weak plications. Radial ornament multicostellate 
with 28-30 primary ribs and about 3-5 costellae per mm along the 
anterior margin of full grown specimens. Growth lines fine, ridge-like, 
crowded, evenly spaced. 

Ventral interior with teeth and long, diverging dental plates continu- 
ing into ridges bordering laterally slightly elongate suboval muscle 
field. Ventral adductor scars narrow and subtriangular. raised anteriorly 
and somewhat shorter than the strongly impressed elongate suboval 
diductor scars. Ventral mantle canals lemniscate, straight, widely di- 
verging. Dorsal interior with high, triangular brachiophores with slightly 
diverging bases. Fulcral plates variably developed. Cardinal process 
ridge-like with crenulated myophore. Dorsal adductor scars bisected 
by fine median ridge and bordered laterally by subparallel ridges 
starting from the ends of the brachiophore bases. 



Discussion. This species is characterised by a uniplicate ante- 
rior commissure with low dorsal median fold and shallow ventral 
sulcus, which is unusual for Pionodema (Cooper 1956), and can 
be compared only with P. uniplicata Cooper, but differs from that 
species in its more transverse outline and the weak umbonal dorsal 
sulcus reversing into a median fold at about mid-length. The 
Anderken specimens also lack the prominent ridge anterior to the 
ventral muscle field of P. uniplicata (Cooper 1956: pi. 154, figs 
31-32). 



Order PENTAMERIDA Schuchert & Cooper, 1931 
Suborder SYNTROPHIIDINA Ulrich & Cooper, 1936 
Superfamily CAMARELLOIDEA Hall & Clarke, 1894 

Family CAMARELLIDAE Hall & Clarke, 1894 
Genus PARASTROPHINA Schuchert & LeVene, 1929 

TYPE SPECIES. Atrypa haemiplicata Hall, 1 847, from the Trenton 
Limestone (Caradoc), New York, U.S.A. 

Parastrophina iliana sp. nov. 

PI. 13, figs 30-50; Figs 15, 16 

1956 Camerella haemiplicata (Hall) var. rotunda (Winchell & 

Schuchert); Rukavishnikova: 129, pi. 2. figs l,3(no«fig. 

2). 
1975 Parastrophina haemiplicata (Hall); Sapelnikov & 

Rukavishnikova: 25, pi. 1, figs 1-8. 
1986 Parastrophina haemiplicata (Hall): Kolobova & Popov; 

pi. Lfig. 4. 

HOLOTYPE. BC 57557, PI. 13, figs 38^2. conjoined valves 
from Sample 100, Anderkenyn-Akchoku section. 

Material. 52 pairs of conjoined valves, one ventral and 3 
dorsal valves from Samples 100 (=K-98/1970) (BC 56643-46, 
57557), 626 (BC 56634-6), Anderkenyn-Akchoku section; Sam- 
ples, 628 (BC 56633, 56642, 57559), 2538 (BC 56637^1 ), 8217 
(BC 57556), 8256 (BC 57558), 85258, Kujandysai Section; Sam- 
ple 948 (BC 57192-99). Tesik River; Sample 1041a, Burultas 
Valley. 

Description. Shell dorsibiconvex to biconvex, transverse, 
semielliptical in outline, about 90% as long as wide and 75% as 
thick as long. Anterior commissure uniplicate. Ventral valve gen- 
tly convex with maximum thickness somewhat posterior to 
mid- valve. Sulcus originating about 5-6 mm from the umbo, 
deepening anteriorly and terminating in broad, semioval tongue 



PLATE 12 

Figs 1-9 Phragmorthis conciliate! Popov. 1, 2, Sample 1018, area 7 km SW of Karpkuduk well. Kotnak Mountains. CNIGR 10/1 1989. holotype. latex 
cast and dorsal internal mould, x 4. 3, 5-9, Sample 2538, Akchoku Mountain, Kujandysai section; 3, BC 57538, dorsal exterior, x 4; 5-8, BC 57539, 
conjoined valves lateral, dorsal, posterior and ventral views, x 4; 9. CNIGR 12/1 1989, latex cast of dorsal exterior, x 4. 4, Sample. 8230. Buldukbai- 
Akchoku section, west side of Kopalysai, BC 57603, dorsal internal mould, x 5. 

Figs 10-12, 15 Eodalmanella extern Popov. 10, Sample 1018, area 7 km SW of Karpkuduk well, Kotnak Mountains, CNIGR 9/1 1989, ventral exterior, x 
4. 11, Sample 620, Anderkenyn-Akchoku section, BC 57189. dorsal exterior latex cast, x 5. 12, 15. Sample 843. Anderkenyn-Akchoku section: 12. 
CNIGR 5/1 1989, dorsal internal mould, x 4; 15, CNIGR 3/1 1989, ventral internal mould, x 4. 

Figs 13, 14, 16-27 Pionodema opima sp. nov. 13, 14, Sample 8257, Buldukbai-Akchoku section, west side of Kopalysai, BC 57544, internal mould, 
dorsal and ventral views, x 3. 16, Sample 8230, east side of Kopalysai. BC 57803, latex cast of dorsal external mould, x 2.7. 17, 18. Sample 7613, 
Akchoku Mountain, Kujandysai section, BC 57545, holotype, internal mould of conjoined valves, dorsal and ventral views, x 3. 19-24. Sample 818a. 
Burultas valley. BC 57802, conjoined valves, detail of radial ornament, x 8, and lateral, ventral, dorsal, posterior and anterior views, x 4. 25, Sample. 
8229. Buldubai-Akchoku section, west side of Kopalysai. BC 57546. dorsal internal mould, x 3. 26, 27, Sample 8228, east side of Kopalysai: 26, BC 
57548, ventral internal mould, x 3; 27, BC 57547, ventral internal mould, x 2. 

Figs 28-35 Ilistrophina tesikensis gen. et sp. nov.. Sample 948, Tesik River. 28-31, BC 56824, conjoined valves, lateral, anterior, ventral and dorsal views, 
x 4. 32-35, BC 56823, conjoined valves, holotype, dorsal, ventral, lateral and anterior views, x 4. 



UPPER ORDOVICIAN BRACHIOPODS FROM KAZAKHSTAN 



65 




66 



L.E. POPOV, L.R.M. COCKS AND I.F. NIKITIN 




Fig. 15 Photographs of transverse serial sections, x 12. Distance in mm is measured from the posterior tip of ventral beak. Dorsal valve uppermost. 1-2, 
Placotriplesia spissa sp.nov., BC 57605, Sample 628, Kujandysai section; 1, 1.5 mm; 2, 1.8 mm. 3, 4, 9-14, Ilistrophina tesikensis gen. et sp. nov., 
Sample 948, Tesik River, 3,4, BC 57606, 0.3 and 0.4 mm; 9-14, BC 57607, 0.4, 0.6, 0.8, 1.0 and 1.4 mm; 5-7, Parastrophina iliana sp. nov., BC 56560, 
Sample 948, Tesik River; 0.6, 0.8 and 1.2 mm; 8, Parastrophina plena Sapelnikov & Rukavishnikova, BC 57564. Sample 948. Tesik River, 1.0 mm. 



about 66% valve width. Dorsal valve strongly convex with maxi- 
mum thickness at mid-valve or slightly anteriorly. Umbonal area 
strongly swollen. Broad median fold originating near mid-valve. 
Radial ornament of coarse rounded-angular ribs originating anterior 
to mid- valve in mature specimens with 1-3 ribs in the sulcus, 2-4- on 
the median fold and up to 6 on the flanks of both valves. 

Ventral interior with small teeth and narrow spondylium on a low 
median septum extending anteriorly to mid-valve. Dorsal interior 
with narrow cruralium on the low median septum. Alate plates small, 
projecting anteriorly as short brachial processes. Inner plates narrow, 
gently curved. Outer plates nearly straight in cross-section, converg- 
ing towards a thin median septum. 

Measurements. (47 1/1 2375) conjoined valves, L= 14.0, W=l 6.9, 



T=6.3, Sw=8.9; (474/12375) conjoined valves, L=20.4, W=21.0, 
T=11.2, Sw=8.7; (475/12375) conjoined valves, L=22.9, W=24.5, 
T=10.2, Sw=12.8; (476/12375) conjoined valves, L=16.5, W=19.8, 
T=6.5, Sw=11.2; (479/12375) conjoined valves, L=10.5, T=6.7, 
Sw=6.7. 

Discussion. The Kazakh shells are comparable to Parastrophina 
haemiplicata (Hall), and in particular with specimens described and 
illustrated by Cooper (1956: 606, pi. 106, figs 33^14; pi. 117, figs 
19-27) from the lower Martinsburg Formation of Virginia, in outline 
and profile of both valves, as well as in the characters of the radial 
ornament, dorsal median fold and ventral sulcus, but they differ in 
having more conspicuous ribbing on the flanks of mature shells, 
which may possess up to 7 ribs; however, the number of ribs is 



Table 23 Measurements of complete shells of Parastrophina iliana sp. nov.. Sample 948, Tesik river. 





L 


W 


T 


Ld 


Sw 


St 


LAV 


T/L 


Ld/W 


Sw/W 


N 


11 


10 


10 


11 


10 


10 


10 


10 


10 


9 


X 


8.2 


9.1 


5.8 


8.1 


6.1 


3.6 


89.2% 


74.0% 


87.7% 


65.6% 


S 


2.05 


2.87 


1.16 


2.10 


1.65 


1.52 


6.3 


10.4 


5.2 


6.5 


MIN 


5.5 


5.6 


4.2 


5.3 


3.2 


1.6 


80% 


62% 


79% 


57% 


MAX 


11.9 


14.8 


7.4 


11.7 


8.5 


6.6 


98% 


95% 


95% 


78% 



UPPER ORDOVICIAN BRACHIOPODS FROM KAZAKHSTAN 



67 




5mm 

i i i i i i 



-k^' 



1.2 









I L 



5 mm 

j i i 



Fig. 16 Transverse serial sections from Sample 948, Tesik River. A, Parastrophina iliana sp. nov., BC 57560; B, Parastrophina plena Sapelnikov & 
Rukavishnikova, BC 57564. Distance in mm is measured from the posterior tip of ventral beak. Dorsal valve uppermost. 



variable in the studied samples. There is also a strong tendency to 
asymmetry in the anterior commissure of the Kazakh shells. How- 
ever, as mentioned by Cooper, his specimens were somewhat different 
from Hall's types and the species needs more substantial revision. 
Specimens from the Anderken Formation described by 
Rukavishnikova (1956) as Camerella haemiplicata (Hall) var. ro- 
tunda seem likely to represent a mixture of several taxa. In particular, 
the specimens illustrated on her pi. 2, fig. 2 may belong to Liostrophia, 
but others appear to be conspecific with ours. It is possible that the 
Kazakh shells are conspecific with the specimens described as 
Parastrophina haemiplicata by Fu (1982: 129, pi. 36, fig. 16) from 
the Jinhe Formation (Caradoc) of northwest China. The only 
illustrated specimen is similar to some of the juvenile Kazakh shells 
in ribbing and in the lateral profile of both valves, but it is impossible 
to estimate the limits of morphological variation in the Chinese 
population of Parastrophina from the published illustrations and 
description. There is remarkable general similarity between the 
brachiopod assemblage from the Jinhe Formation and the fauna from 
the carbonate mud-mounds in the upper Anderken Formation. In 
particular, both assemblages contain distinctive genera such as 



Schizostrophina and Pectenospira (Popov et al. 
otherwise unknown elsewhere. 



1999) which are 



Parastrophina plena Sapelnikov & Rukavishnikova, 1975 

PI. 13, figs 51-58; Figs 15, 16 



1975 



1982 



Parastrophina plena Sapelnikov & Rukavishnikova: 27, pi. 

l.figs 12-14. 

Parastrophina uniplicata Fu:130, pi. 36, fig. 17. 



HOLOTYPE. IGNA 436/1 86 1 , conjoined valves, from the Anderken 
Formation, Sample 302 of Keller (1956), Anderkenyn-Akchoku 
section. 

Material. 102 pairs of conjoined valves, 1 ventral and 2 dorsal 
valves from Samples 100 (=K-98/1970), 8223a (BC 56595), 8223b 
(BC 57248-52), 8226, Anderkenyn-Akchoku section; Samples 8214 
(BC 57216-37, 57563), 8215 (BC 57238-47), west side of Ashchisu 
River; Samples 628 (BC 56598-600), 2536 (BC 56605-10), 2538 
(BC 57562), 8217, 8219, 8256 (BC 56603-4), Kujandysai Section; 
Sample 948 (BC 57200-15), Tesik River. 



Table 24 


Measurements of ventral valves of Par 


astrophina plena 


Sapelnikov 


and Rukavishnikova, Sample 


948. Tesik river. 








L 


W 


T 


Ld 


Sw 


St LAV 


T/L 


LdAV 


SwAV 


N 


34 


34 


34 


34 


31 


33 34 


34 


34 


31 


X 


6.6 


7.1 


4.8 


6.7 


4.3 


3.1 93.6% 


72.7% 


94.3% 


60.7% 


s 


0.74 


0.95 


0.91 


0.75 


0.55 


0.82 5.7 


8.7 


5.5 


5.3 


MIN 


5.5 


5.8 


3 


5.5 


3.4 


1 .4 83.0% 


52.6% 


83.6% 


52.9% 


MAX 


9 


9.7 


7.5 


9 


5.4 


4.8 107.5% 


87.7% 


107.5% 


76.8% 



68 



L.E. POPOV, L.R.M. COCKS AND I.F. NIKITIN 

i 





F 47 ^48 49 




53 ^piPSP- j£. 
59 







^1^ 62 ^f 



58 





63 




61 67 



Soy j« 




68 



71 




64 ^*!^ 65 66 



69 





l^tv 



70 



72 



M?,?^ 



UPPER ORDOVICIAN BRACHIOPODS FROM KAZAKHSTAN 



69 



Description. Shell smooth, dorsibiconvex, subpentagonal in out- 
line, about 70% as thick as long and 95% as long as wide. Anterior 
commisure sulciplicate. Ventral valve gently convex in lateral profile 
with maximum thickness about one-third valve length from the 
small, slightly erect pointed beak posterior to the hinge line. Sulcus 
about 60% as wide as the valve, originating in the umbonal area at 
about 4-5 mm from the beak, strongly deepening anteriorly and 
divided medianly by high angular costa originating near mid-valve. 
Dorsal valve strongly and evenly convex with maximum thickness at 
mid-valve with median fold originating slightly anterior to mid- 
valve and bearing two strong angular ribs. 

Ventral interior with small teeth and deep, narrow spondylium on 
the low median septum. Dorsal interior with narrow cruralium raised 
anteriorly on a low median septum extending anteriorly not more 
than 1 .5 mm in adults. Inner plates small, thickened, slightly curved 
in cross-section. Brachial processes short. 

Measurements. (47 1/1 2375) conjoined valves, L=14.0,W=1 6.9, 
T=6.3, Sw=8.9; (474/12375) conjoined valves, L=20.4, W=21.0, 
T=11.2, Sw=8.7; (475/12375) conjoined valves, L=22.9, W=24.5, 
T=10.2, Sw=12.8; (476/12375) conjoined valves, L=16.5, W=19.8, 
T=6.5, Sw=11.2; (479/12375) conjoined valves, L=10.5, T=6.7, 
Sw=6.7. 

Discussion. This species usually occurs in the upper Anderken 
Formation together with Parastrophina iliana, but can be easily 
distinguished in being half the size, in the smooth shell with a single 
costa in the sulcus and two in the fold of mature specimens, and in the 
sulciplicate anterior commissure. 

Parastrophina plena closely resembles Parastrophina uniplicata 
Fu, 1982 from the Jinhe Formation (Caradoc) of northwest China in 
its smooth shell, sulciplicate anterior commissure, and in the 
characters of the dorsal fold and ventral sulcus. They may be 
conspecific. 



Genus IL1STROPHINA Gen.nov. 

TYPE SPECIES. Uistrophina tesikensis sp.nov. 

Diagnosis. Shell smooth, ventribiconvex with well-developed 
dorsal median fold and ventral sulcus; ventral interior with 
spondylium sessile posteriorly, raised anteriorly on low median 
septum; dorsal interior with cruralium supported by high median 
septum; well defined alate plates and long brachial processes. 

DISCUSSION. Externally Uistrophina is most similar to Eostrophina 
(Zhan & Rong 1995) but differs in having a narrow spondylium 
sessile posteriorly, raised anteriorly on a low median septum and 
well developed alate plates. It differs from Liostrophia in having 
strongly raised cruralium supported by a median septum instead of a 
sessile cruralium or separated outer plates; however, present know- 
ledge of the interior of type species of the former remains inadequate. 
Uistrophina resembles Parastrophina and Parastrophinella in the 
interior morphology of both valves, but differs in having a smooth 
shell and a sessile spondylium posteriorly. Jin and Copper (1997) 
demonstrated that in Parastrophinella the cruralium is supported by 
the median septum along its entire length, but, in contrast to 
Uistrophina, the dorsal median septum in Parastrophinella is very 
low and buried within secondary shell in the apical area. 

Uistrophina tesikensis sp. nov. 

PI. 12. figs 28-35; Figs 15, 17 

Etymology. After the type locality. 

HOLOTYPE. BC 56823, PI. 12, figs 32-35, conjoined valves, from 
Sample 948, Tesik River. 

Material. 14 pairs of conjoined valves (including BC 56823-24) 
and one dorsal valve, all from Sample 948, Tesik River. 

Description. Shell smooth, dorsibiconvex, subcircular in outline, 



Table 25 Measurements of ventral valves of Uistrophina tesikensis sp. nov.. Sample 948, Tesik river. 





L 


W 


T 


Ld 


Sw 


St 


LAV 


T/L 


LdAV 


Sw/W 


N 


9 


9 


9 


9 


9 


8 


9 


9 


9 


9 


X 


7.5 


8.2 


5.1 


7.5 


5.7 


3.9 


92.3% 


68.2% 


92.2% 


70.2% 


s 


1.01 


1.46 


0.82 


0.98 


0.96 


0.94 


4.7 


6.6 


4.9 


5.4 


MEM 


6.2 


6.4 


4.2 


6.2 


4.6 


2.4 


83.8% 


58.3% 


82.9% 


63.9% 


MAX 


9.8 


11.7 


6.8 


9.7 


7.8 


5 


97.5% 


79.4% 


97.5% 


80.7% 





PLATE 13 

Figs 1-29 Liostrophia pravula sp. nov., Akchoku Mountain, Kujandysai section. 1-15, Sample 8256; 1-5, BC 57550, conjoined valves, ventral, dorsal, 
anterior, posterior and lateral views, x 2; 6-10, BC 57551, conjoined valves, dorsal, ventral, anterior, posterior and lateral views, x 2; 11-15, BC 57552, 
conjoined valves, dorsal , ventral, lateral, posterior and anterior views, x 2. 16-29, Sample 2538; 16-20, BC 57553, holotype, conjoined valves, dorsal, 
ventral, lateral, anterior and posterior views, x 2; 21-24, BC 57554, conjoined valves, ventral, dorsal, anterior and lateral views, x 2; 25-29, BC 57555, 
conjoined valves, ventral, dorsal, posterior, lateral and anterior views, x 2. 

Figs 30-50 Parastrophina iliana sp.nov. 30-33, Sample 8217, Kujandysai section, BC 57556, conjoined valves, dorsal, anterior, lateral and ventral views, 
x 2. 34-37, Sample 2538, Kujandysai section, CNIGR 4/12361, conjoined valves, lateral, dorsal, ventral and posterior views, x 2. 38-42, Sample 100, 
Anderkenyn-Akchoku section, BC 57557, holotype, conjoined valves, dorsal, ventral, posterior, anterior and lateral views, x 2. 43-46, Sample 8256, 
Kujandysai section, BC 57558, conjoined valves, dorsal, lateral, ventral and posterior views, x 2. 47-50, Sample 628, Kujandysai section, BC 57559, 
conjoined valves, dorsal, lateral, ventral and anterior views, x 2. 

Figs 51-58 Parastrophina plena Sapelnikov & Rukavishnikova, 1975. 51-54, Sample 2538. Kujandysai section, BC 57562, conjoined valves, lateral, 
dorsal, ventral and anterior views, x 2. 55-58, Sample 8214, Anderkenyn-Akchoku section, BC 57563, conjoined valves, anterior, lateral, ventral and 
dorsal views, x 2. 

Figs 59-74 Plectosyntrophia unicostata sp. nov., Anderkenyn-Akchoku section. 59-62, 67-70, Sample 626; 59-62, BC 57568, conjoined valves, dorsal, 
posterior, lateral, and ventral views, x 2; 67-70, BC 57570, conjoined valves, holotype, lateral, ventral, dorsal and posterior views, x 2. 63-66, 71-74, 
Sample 100; 63-66, BC 57569, conjoined valves, lateral, anterior, dorsal and posterior views, x 2; 71-74, BC 57571, conjoined valves, lateral, dorsal 
ventral and anterior views, x 2. 

Figs 75-77 Didymelasma cf. transversa Fu, Sample 2538. Kujandysai section. BC 57573, conjoined valves, dorsal, lateral and ventral views, x 2. 



70 



L.E. POPOV, L.R.M. COCKS AND I.F. NIKITIN 



&&OTW" 






Fig. 17 Transverse serial sections of Ilistrophina tesikensis gen. et sp. nov.. BC 57606. Sample 948. Tesik River. Distance in mm is measured from the 
posterior tip of ventral beak (dorsal valve uppermost), also an axial diagram showing the plates. 



about 65% as high as long and 90% as long as wide with maximum 
width at mid-length. Anterior commissure strongly uniplicate. Ven- 
tral valve gently convex in lateral profile with maximum thickness 
somewhat posterior to mid-length. Beak small, curved towards the 
hinge line. Sulcus originating about 4-5 mm from the umbo, strongly 
deepening anteriorly and terminating with a high semioval tongue 
about 70% valve width. Dorsal valve with moderately convex lateral 
profile. Beak slightly swollen, curved. Dorsal median fold originat- 
ing anterior to mid-length, semioval in cross-section. 

Ventral interior with small teeth and narrow spondylium, bell- 
shaped in cross-section, sessile posteriorly and raised anteriorly on a 
low, thick median septum extending to mid-valve length. Dorsal 
interior with cruralium supported by high median septum along its 
entire length. Inner plates present. 

Measurements, conjoined valves, Lv=9.8, W=11.7. T=6.8, 
Sw=7.8, St=4.0; conjoined valves, Lv=7.8, W=8.2, T=5.7; Sw=5.5, 
St=4.7. 

Discussion. This species is externally comparable to Eostrophina 
uniplicata from the Middle Ashgill Xiazhen Formation of South 
China, but can be easily distinguished in its smaller size, which does 
not exceed 10 mm in length in the largest specimens, and in having 
a dorsal median fold evenly rounded in cross-section and originating 
anteriorly to the mid-valve. 



Genus LIOSTROPHIA Cooper & Kindle. 1936 

Type SPECIES. Liostrophia glabra Cooper & Kindle. 1936, from 
the Ashgill of Canada. 

Liostrophia pravula sp. nov. PI. 13, figs 1-29; Fig. 18 

Etymology. After pravus, Latin - bowed. 

HOLOTYPE. BC 57553, pi. 13, figs 16-20, conjoined valves, from 
the Anderken Formation, Sample 2538, Akchoku Mountain, 
Kuyandysai section. 



Material. 30 pairs of conjoined valves, one ventral and 7 dorsal 
valves, from Samples 100 (=K-98/1970) (BC 57253-56), 626 (BC 
57263-65), 8223, Anderkenyn- Akchoku section; Samples 2538 (BC 
57266-69, 57553-55), 8256 (BC 57550-52), Kuyandysai section; 
Sample 948 (BC 57257-62), Tesik River. 

Description. Shell slightly dorsibiconvex, transverse, subcircular 
in outline, about 5 1 % as thick as long and 90% as long as wide, with 
maximum width at mid-length. Anterior commissure gently 
uniplicate. Ventral valve gently convex in lateral profile, with maxi- 
mum thickness somewhat posterior to mid-length. Small, pointed 
beak, slightly erect and posterior to hinge line. Broad shallow sulcus, 
originating near the anterior margin, usually asymmetrical in cross- 
section. Dorsal valve moderately and evenly convex in profile with 
slightly swollen, curved beak. Dorsal median fold usually absent but 
weakly developed anteriorly in mature specimens. Shell surface 
smooth with one or two ribs in the fold and sulcus near the anterior 
margin of the largest specimens. 

Ventral interior with teeth and spondylium supported anteriorly 
by very low septum enclosed posteriorly by a secondary shell 
thickening (Fig. 18). Dorsal interior with sessile cruralium formed 
by high outer plates joined at the valve floor. Alate plates small, 
appearing at some distance from the umbo. Inner plates small, 
slightly thickened, strongly curved in cross-section with the outer 
parts located almost within comissural plane. Brachial processes 
thin and relatively long. 

Measurements. (518/12375) conjoined valves, L=7.6, W=7.9, 

Table 26 Measurements of ventral valves of Liostrophia pravula sp. nov.. 
Sample 2538. Kujandysai section. 





L 


W 


T 


Sw 


St 


LAV 


T/L 


SwAV 


N 


11 


11 


11 


10 


10 


9 


9 


9 


X 


5.7 


6.3 


2.9 


4.1 


1.5 


89.7% 


50.9% 


61.8% 


s 


1.45 


1.56 


1.05 


0.75 


0.18 


7.6 


6.3 


3.7 


MIN 


3.3 


3.5 


1.3 


3.2 


1.2 


81.5% 


42.6% 


57.1% 


MAX 


8.2 


9.2 


5.1 


5.7 


1.8 


103.8% 


62.2% 


70.2% 



UPPER ORDOVICIAN BRACHIOPODS FROM KAZAKHSTAN 



71 



sessile crurallum 




1(0-2) 2(0.3) 3(0.4) 4(05) ^Y(0. 6) 




11 (1.5) 




median septum 



Fig. 18 Transverse serial sections of Liostrophia pravula sp.nov.. Sample 2538, Kujandysai section. Distance in mm is measured from the posterior tip of 
ventral beak. Dorsal valve uppermost. Also lateral view to show section positions and schematic reconstruction. 



T=3.4; (519/12375) conjoined valves, L=5.2, W=5.7, T=2.0; (520/ 
12375) conjoined valves, L=7.9, W=7.9, T=3.2; (521/12375) con- 
joined valves, L=8.7, W=8.2, T=4. 1 ; (522/12375) conjoined valves, 
L=5.8, T=6.3, Sw=3.7.1; (523/12375) conjoined valves, L=10.6, 
T=10.5, Sw=4.7. 

DISCUSSION. This species differs from Liostrophia glabra Cooper 
& Kindle in having a subcircular outline, weak ventral sulcus and in 
the absence of a dorsal median fold. The characters of the dorsal 
interior in the type species remain inadequately known. In particular 
it is unclear from the existing illustrations whether or not it has a 
sessile cruralium or if it is supported anteriorly by a very short 
septum. In external morphology, particularly in the smooth, rounded 
shell with an uniplicate anterior commissure but without a distinct 
dorsal median fold, Liostrophia pravula resembles Psilocamera 
planisulcata Fu, 1982 from the the Jinhe Formation (Caradoc) of 
north-west China. However, in the single transverse section provided 
by Fu (1982, text-fig. 1 8A) the outer plates appear to be completely 
separate and there are no alate plates or inner plates illustrated. 
Liostrophia pravula differs from juvenile specimens of llistrophina 



Table 27 Measurements of ventral valves of Plectosyntrophia unicostata 
sp. nov., samples 100 and 626 from Anderkenyn-Akchoku section. 





L 


W 


T 


St 


Sw 


LAV 


T/L 


Sw/W 


N 


6 


6 


6 


6 


6 


6 


6 


6 


X 


9.5 


10.7 


7.2 


4.0 


5.8 


89.4% 


74.1% 


156.2% 


s 


1.37 


2.10 


2.77 


1.87 


1.82 


4.6 


18.3 


28.4 


MIN 


7.8 


8.4 


4.6 


2.2 


3.5 


82.9% 


59.0% 


119.6% 


MAX 


11 


13.2 


11.2 


6.5 


7.9 


92.9% 


101.8% 


177.1% 



tesikensis not only in its larger size and sessile cruralium (which 
hardly exceeds half the maximum length), but also in the absence of 
a dorsal median fold. A ventral sulcus is present in llistrophina 
tesikensis when specimens are 4-5 mm long, whereas in Liostrophia 
pravula it is visible only in mature specimens which exceed the 
average shell size of about 6 mm. 



Subfamily ANASTROPHIINAE Nikiforova, 1960 
Genus PLECTOSYNTROPHIA Fu, 1982 



TYPE SPECIES. Plectosyntrophia qilianshanensis Fu, 1982. 
the Yingou Group (Middle Ordovician) of North China. 



from 



Plectosyntrophia unicostata sp. nov. 

PI. 13, figs 59-74; Fig. 19 

Holotype. BC 57570, PI. 13, figs 67-70, conjoined valves from 
Sample 626, Anderkenyn-Akchoku section. 

Material. Seven pairs of conjoined valves and one dorsal valve 
from Samples 100 (=K-98/1970) (BC 57569, 71), 620 (BC57360), 
626 (BC 57271-73, 57568, 70, 72), 8214 (BC 57361), Anderkenyn- 
Akchoku section; Samples 2538, 8217 (BC 57362), Kujandysai 
Section. 

DESCRIPTION. Shell subequally biconvex, transverse, subpent- 
agonal in outline, about 75% as thick as long and about 89% as long 
as wide with maximum width slightly anterior to mid-length. Ante- 
rior commissure uniplicate. Ventral valve moderately convex with 
curved beak slightly raised above a narrow triangular, apsacline 
interarea. Ventral sulcus narrow and shallow, but with steep lateral 



72 



L.E. POPOV, L.R.M. COCKS AND I.F. NIKITIN 




Fig. 19 Photographs of transverse serial sections of Plectosyntrophia unicostata sp. nov., BC 57572 
12. Distance in mm is measured from the posterior tip of ventral beak. Dorsal valve uppermost. 



from Sample 626, Anderkenyn-Akchoku section: x 



slopes, originating slightly posteriorly to mid- valve, and ending with 
a shallow, narrow, trapezoidal tongue about 40% valve width. Dorsal 
valve strongly convex in lateral profile with maximum thickness 
near mid-length. Beak slightly swollen and strongly curved towards 
the hinge line. A low, flattened median fold originates from the 
umbo. Radial ornament mainly costate, with occasional bifurcating 
ribs and with 1 primary rib in the sulcus, 2 primary ribs in the median 
fold and 6-9 on the lateral slopes of the valves. In some specimens 
one or two small secondary costellae originate in the median fold and 
dorsal sulcus between the umbo and mid-length. Ventral interior 
with strong teeth; bell-shaped spondylium in transverse section, 
sessile posteriorly and raised anteriorly on a low median septum 
partly covered by secondary shell (Fig. 19). Dorsal interior with 
narrow cruralium on a high median septum extending anteriorly up 
to 3 mm in adults. Inner plates narrow, curved; alate plates narrow, 
bordered laterally by a pair of high subparallel muscle bounding 
ridges. 

Measurements. (CNIGR 471/12375) conjoined valves, L=14.0, 
W=16.9, T=6.3, Sw=8.9; (474/12375) conjoined valves, L=20.4, 
W=21.0, T=l 1.2, Sw=8.7; (475/12375) conjoined valves, L=22.9, 
W=24.5, T=10.2, Sw=12.8; (476/12375) conjoined valves, L=16.5, 



W=19.8, T=6.5, Sw=11.2; (479/12375) conjoined valves, L=10.5, 
T=6.7, Sw=6.7. 

Discussion. The generic attribution of unicostata is tentative 
because the interior of Plectostrophia qilianshanensis is inadequately 
known. In particular, there is no record of the presence of alate plates 
in the type species, but some illustrations provided in the original 
description (Fu 1982, text-fig. 17) suggest their presence. The 
characters of the ventral interior, in particular the presence of a 
sessile spondylium slightly raised near its anterior margin on a short 
septum, also needs confirmation, because Fu's illustrations are sche- 
matic and it could be a preservational pattern. In the Kazakh species 
alate plates are well defined, whereas the sessile spondylium is 
present only in the earliest ontogenetic stages and is characterised by 
a spondylium supported by a low median septum partly covered by 
secondary shell. Plectosyntrophia qilianshanensis has 3 ribs in the 
ventral sulcus and 4 in the dorsal fold, which makes it more similar 
to Eoanastrophia kurdaica than to our species. 

P. unicostata differs from Eoanastrophia kurdaica Sapelnikov & 
Rukavishnikova ( 1 975 ) from the Keskentas Formation (Caradoc), of 
the Kendyktas Range, south Kazakhstan, in having a more pro- 
nounced dorsal median fold with two primary ribs and a ventral 



UPPER ORDOVICIAN BRACHIOPODS FROM KAZAKHSTAN 



73 



sulcus with a single accentuated primary rib originating in the 
umbonal area. The coarsely ribbed radial ornament, well defined 
dorsal median fold and ventral sulcus resembles numerous species of 
Plectocamara described by Cooper (1956) from the Caradoc of 
North America, but there is no evidence of the presence of alate 
plates in the former genus. 



Family PARALELLELASMATIDAE Cooper, 1956 
Genus SCHIZOSTROPHINA Fu, 1982 

Type species. Schizostrophina margarita Fu, 1 982, from the Jinhe 
Formation (Caradoc), northwest China. 

DIAGNOSIS (emended). Shell equally biconvex with parasulcate 
anterior commissure; shallow ventral median sulcus and dorsal 
median fold originating in umbonal area; radial ornament of variably 
developed coarse angular ribs in posterior half of the shell; ventral 
interior with delicate teeth and spondylium supported posteriorly 
and free anteriorly; dorsal interior with separated outer plates, slightly 
diverging distally, and well-defined brachial processes. 

Discussion. Schizostrophina was erected originally by Fu ( 1982) 
without proper illustrations and detailed description of the interior in 
the type species. However, the very distinctive exterior morphology 



of the shell, which is unique in late Ordovician syntrophiidines, 
leaves no doubt that the Kazakh shells from the Anderken Formation 
belong to the same genus and species. Their internal morphology 
confirms the original assignment of the genus to the Paralellel- 
asmatidae and suggests a close affinity to Paralellelasma, but 
Schizostrophina lacks the radial capillae and the characteristic trun- 
cated margins of the ribs along the commissure of the former genus. 
Another difference is the parasulcate anterior commissure. 

Schizostrophina margarita Fu, 1982 

PI. 14, figs 2-27; Fig. 20 

1982 Schizostrophina margarita Fu: 132, pi. 37, fig. 5. 
1982 Schizostrophina shaanxiensis Fu: 133, pi. 37, fig. 6. 

Material. 26 pairs of conjoined valves, 6 ventral and 4 dorsal 
valves from Samples 100 (=K-98/1970) (BC 57275-7), 626 (BC 
57279-8 1 ), 8223, Anderkenyn- Akchoku section; Samples 628, 2538 
(BC 57282-92, 57574-78), 8219 (BC 57294), 8220 (BC 57295), 
8256 (BC 57296-9), Kujandysai Section. 

Description. Shell equally biconvex, subpentagonal to 
subtriangular in outline; about as long as wide, with maximum width 
anterior to mid-valve. Anterior commissure parasulcate. Beaks of 
both valves swollen and strongly curved. Ventral valve moderately 



Table 28 Measurements of ventral valves of Schizostrophina margarita Fu, Samples 626, 2538, 8256 from Anderkenyn- Akchoku and Kujandysai sections. 





L 


W 


T 


Sw 


St 


LAV 


T/L 


SwAV 


LdAV 


SwAV 


N 


18 


18 


18 


17 


3 


18 


18 


17 


9 


9 


X 


5.8 


6.2 


4.2 


4.3 


4.3 


94.8% 


71.4% 


71.7% 


92.2% 


70.2% 


s 


1.24 


1.68 


1.25 


1.55 


0.92 


6.0 


9.3% 


14.5 


4.9 


5.4 


MIN 


3.6 


3.7 


2.1 


2.6 


3.2 


79.6% 


54.5% 


45.2% 


82.9% 


63.9% 


MAX 


8.0 


9.8 


6.2 


7.8 


4.8 


102.2% 


90.4% 


100.0% 


97.5% 


80.7% 



discrete outer plates 




G) 

1 (0.1) 



2 (<>-2) 3 (0.3) 



4 (0.4) 



5 (0.6) 



6 (0.8) 




5 mm 



5 mm 



Fig. 20 Transverse serial sections of Schizostrophina margarita Fu, Sample 2538, Kujandysai section. Distance in mm is measured from the posterior tip 
of ventral beak. Dorsal valve uppermost. Also lateral view to show section positions and schematic reconstruction. 



74 



L.E. POPOV, L.R.M. COCKS AND I.F. NIKITIN 



convex in lateral profile with maximum thickness slightly posterior 
to mid-valve. Ventral sulcus originating near the umbo, broad and 
shallow with low trapezoidal tongue slightly inclined anteriorly. 
Lateral slopes steep, slightly convex in cross-section. Dorsal valve 
with moderately convex lateral profile strongly curved posteriorly. 
Median fold shallow, slightly rounded in transverse section, origi- 
nating near the beak, flanked by two strong plications. Radial 
ornament with angular ribs originating near mid-length, 1-3 ribs in 
the sulcus and 2-4 in the median fold. Pair of ribs occasionally on the 
lateral slopes of both valves. Ventral interior with deep, narrow 
spondylium about one-sixth valve length, supported posteriorly by 
high median septum, free anteriorly. Dorsal interior with separated 
outer plates, slightly diverging distally, short subtriangular brachial 
plates and well-defined brachial processes. 

Measurements, conjoined valves, L=7.9, W=8.6,T=5.9, Sw=6.9; 
conjoined valves, L=9.8, W= 11.2, T=6.7, Sw=8.2; conjoined valves, 
L=7.8, W=8.2, T=6.2, Sw=6.8; conjoined valves, L=6.8, W=7.2, 
T=6.2, Sw=5.4; conjoined valves, L=5.7, W=5.8, T=3.4, Sw=4.6. 

Discussion. Schizostrophina margarita, the type species of the 
genus, came from the same unit and locality, the Jinhe Formation 
(Lower Caradoc) of north-west China, as S. shaanxiensis and differs 
from the latter in having a single poorly defined rib in the ventral 
sulcus and smooth lateral sides of the valve. The Kazakh shells 
demonstrate a strong variability in the number and characters of 
radial ornament with growth. The small shells (about 5-6 mm long) 
usually lack ribs on the lateral sides of the shell, and the radial 
ornament in the ventral sulcus and dorsal fold is poorly developed or 
absent (PI. 14, figs 22-24), whereas mature specimens possess a 
radial ornament closely comparable to one of the specimens referred 
by Fu (1982) to S. shaanxiensis. This suggests that all the shells 
described by Fu represent a single species, which should be termed 
S. margarita. Since there appear to be no constant morphological 
differences between the Kazakh and Chinese specimens of 
Schizostrophina. they are regarded here as conspecific. 



2538, Kujandysai Section; one dorsal internal mould (BC 57366) 
from Sample 1018, 7 km southwest of Karpkuduk well, Kotnak 
Mountains. 

Description. Shell smooth, subequally biconvex, transverse to 
suboval in outline with uniplicate posterior commissure. Ventral 
valve lateral profile moderately convex with maximum thickness 
about one-third valve length. Ventral beak swollen and slightly 
curved. Sulcus originating near mid-length, strongly deepening 
towards the anterior margin, with low median rib. Tongue 
semielliptical. Dorsal valve moderately convex in lateral profile with 
maximum thickness at about two-thirds valve length. Median fold 
originating somewhat posteriorly to mid-valve, clearly separated 
from the slightly convex lateral sides of the valve. Ventral interior 
unknown, except for median septum, possibly supporting 
spondylium. Dorsal interior with separated, long, subparallel outer 
plates. 

Measurements, conjoined valves, L=13.4, T=8.2. 

Discussion. The Kazakh specimens closely resemble Didymel- 
asma transversa Fu, 1982, from the Caradoc Jinhe Formation of 
northwest China, in the external features of their smooth shells, 
including the shape of the dorsal median fold and ventral sulcus, but 
our material is insufficient to make a precise attribution to this 
species. Fu (1982, text-fig. 23) also demonstrated the presence of 
separated, subparallel outer plates in the Chinese shells, which 
supports their assignation to Didymelasma. 



Order RHYNCHONELLIDA Kuhn, 1949 

Superfamily RHYNCHOTREMATOIDEA Schuchert, 1913 

Family RHYNCHOTREMATIDAE Schuchert, 1913 

Genus RHYNCHOTREMA Hall, 1860 

TYPE SPECIES. Atrypa inerebescens Hall, 1847, from the Trenton 
Formation (Caradoc), New York, U.S.A. 



Genus DIDYMELASMA Cooper, 1956 

TYPE SPECIES. Didymelasma longicrurum Cooper, 1956 from the 
Lebanon Formation (Caradoc), Tennessee, U.S.A. 

Didymelasma cf. transversa Fu, 1982 

PI. 13, figs 75-77, PI. 14, fig. 1 



Rhynchotrema akchokense sp. nov. PI. 14, figs 28^42 

Etymology. After Akchoku Mountain on the east side of 
Kopalysai. 

Holotype. BC 57579, PI. 14, figs 28-32, conjoined valves, from 
the Anderken Formation, Sample 626, Anderkenyn-Akchoku sec- 
tion. 



Material. One pair of conjoined valves, BC 57573, from Sample Material. 11 conjoined valves, five ventral and six dorsal valves 



PLATE 14 

Fig. 1 Didymelasma cf. transversa Fu, Sample 1018. area 7 km SW of Karpkuduk well, Kotnak Mountains, BC 57366, dorsal internal mould, x 2. 
Figs 2-27 Schizostrophina margarita Fu, Sample 2538, Kujandysai section. 2—6, BC 57574, conjoined valves, dorsal, ventral, posterior, lateral and 

anterior views, x 3. 7-9, 11, 14, BC 57575, conjoined valves, anterior, dorsal, ventral, lateral and posterior views, x 3. 10, 12, 15-17. BC 57576, 

conjoined valves, anterior, posterior, dorsal, lateral and ventral views, x 3. 13, CNIGR 5/12361, dorsal exterior, x 3. 18-22, BC 57577, conjoined valves, 

lateral , dorsal, ventral, anterior and posterior views, x 3. 23-27, BC 57578. conjoined valves, ventral , dorsal, lateral, posterior and anterior views, x 3. 
Figs 28-42 Rhynchotrema akchokense sp. nov. 28-32, 38-41, Sample 626, Anderkenyn-Akchoku section; 28-32, BC 57579, conjoined valves, holotype. 

ventral, lateral, dorsal, posterior and anterior views, x 2; 38-41, BC 57582, conjoined valves, dorsal, ventral, anterior and lateral views, x 3. 33-36, 

Sample 2538, Kujandysai section, BC 57580, conjoined valves, anterior, dorsal, ventral and lateral views, x 2. 37, 42. Sample 843. Anderkenyn-Akchoku 

section; 37, BC 57581, ventral internal mould, x 3: 42, BC 57583, dorsal internal mould, x 5. 
Figs 43-46 Pectenospira pectenata Popov, Nikitin & Sokiran, Sample 948, Tesik River, BC 57320, dorsal, anterior, ventral and lateral views of conjoined 

valves, x 5.5. 
Figs 47-57 Nikolaispira guttula sp.nov. 47-53, Sample 2538. Kujandysai section; 47-49, BC 57584, conjoined valves, anterior, dorsal and ventral views, 

x 3; 50-53. BC 57585. conjoined valves, holotype. ventral, anterior, lateral and dorsal views, x 3. 54-57, Sample 8221, Anderkenyn-Akchoku section, 

BC56770, conjoined valves, anterior, ventral, dorsal and lateral views, x 4. 
Figs 58-61 Kellerella misiusi Popov, Nikitin & Sokiran, Sample 8214, Anderkenyn-Akchoku section. BC 56773, conjoined valves, anterior, ventral, 

lateral and dorsal views, x 3. 



UPPER ORDOVICIAN BRACHIOPODS FROM KAZAKHSTAN 



75 




76 



L.E. POPOV, L.R.M. COCKS AND I.F. NIKITIN 



from Samples 100 (=K-98/1970) (BC 56656-9), 626, 843 (BC 
5758 1-83), Anderkenyn-Akchoku section; Samples 628 (BC 5666 1 ), 
2538 (BC 57580), 7613 (BC 56649-55), Kujandysai Section; Sam- 
ple 948 (BC 5731 1-2), Tesik River. 

Description. Shell dorsibiconvex to biconvex, slightly transverse, 
about 73% as thick as long and about 75% as long as wide with 
maximum width at mid-length. Anterior commissure uniplicate. 
Ventral valve moderately convex in lateral profile with maximum 
thickness at quarter valve length from the slightly curved, pointed 
beak. Delthyrium open, narrow triangular. Ventral sulcus originating 
2-3 mm from the umbo, very shallow posterior to mid-valve, but 
deepening anteriorly and terminating in low, trapezoidal tongue 
about 79% valve width. Dorsal valve moderately and evenly convex 
in lateral profile with shallow umbonal sulcus inverting into a low 
flattened median fold with steep lateral slopes. Radial ornament of 
coarse angular ribs, usually with 4 ribs in the dorsal median fold, 3 
ribs in the sulcus and 6-8 on flanks of both valves. 

Ventral interior with cyrtomatodont teeth and short, thin dental 
plates flaring close to the sides of the valve. A pedicle base impres- 
sion occupies the floor of the delthyrial cavity; large, weakly 
impressed ventral muscle field with small, lanceolate adductor scars 
completely surrounded by diductor scars anteriorly. Dorsal interior 
with disjunct hinge plate and narrow cruralium bearing a simple, 
ridge-like cardinal process and long, high median ridge extending 
anteriorly to mid-valve. Adjuster scars weakly impressed with pos- 
terior and anterior pair of about equal size, separated by fine, oblique 
transmuscle ridges. 

Measurements. (471/1 2375) conjoined valves, L=14.0,W=16.9, 
T=6.3, Sw=8.9; (474/12375) conjoined valves, L=20.4, W=21.0, 
T=11.2, Sw=8.7; (475/12375) conjoined valves, L=22.9, W=24.5, 
T=10.2, Sw=12.8; (476/12375) conjoined valves, L=16.5, W=19.8, 
T=6.5, Sw=11.2; (479/12375) conjoined valves, L=10.5, T=6.7, 
Sw=6.7. 

Discussion. Rhynchonellides are widespread in the mid and late 
Ordovician shallow-water benthic assemblages of Kazakhstan, but 
with few exceptions are represented exclusively by ancystrorhynchids 
and oligorhynchids (Nikiforova & Popov 1981; Nikitin & Popov in 
Klenina et al. 1 984). All other Kazakh rhynchonellide species previ- 
ously described as Rhynchotrema by Rukavishnikova (1956) and 
Klenina (in Klenina et o/. 1 984) belong in reality to the 
ancystrorhynchid Altaethyrella or atrypides related to Nalivkinia 
(Popov et al. 2000). This species represents the earliest known 
record of rhynchonellides with the cruralium supported by the dorsal 
median septum in Kazakstan. Externally it is similar to Rostricellula 
sarysuica Nikitin & Popov (Nikitin et al. 1996) from the Upper 
Caradoc to Lower Ashgill Dulankara Regional Stage of the northern 
Betpak-Dala Desert, Kazakhstan, but differs in the less convex 
lateral profile of the dorsal valve, with maximum height at mid- 
length, the relatively shallow ventral sulcus, and in the presence of a 
ridge-like cardinal process. 

Rhynchotrema akchokense is similar in radial ornament to two 
Australian species of the genus, R. oepiki Percival, 1991, from the 



Upper Caradoc of New South Wales, and R. bailliei Laurie, 1991, 
from the Caradoc of Tasmania, in having a more transverse shell 
outline, a less convex dorsal valve profile, with maximum height 
near the mid-length, a relatively shallow ventral sulcus and a low 
dorsal median fold. R. bailliei is also characterized by its poorly 
developed cardinal process, which makes its generic assignment 
somewhat questionable, although we refer it to Rhynchotrema. 



Order ATRYPIDA Rzhonsnitskaya, 1960 

Superfamily LISSATRYPOIDEA Twenhofel, 1914 

Family PROTOZYGIDAE Copper, 1986 

Genus PECTENOSPIRA Popov, Nikitin & Sokiran, 1999 

TYPE SPECIES. Pectenospira pectenata Popov, Nikitin & Sokiran, 
1999, from the Anderken Formation, Chu-Ili Range. 

Pectenospira pectenata Popov, Nikitin & Sokiran, 1999 

PI. 14, figs 43^6 

1999 Pectenospira pectenata Popov, Nikitin & Sokiran: 648, pi. 
4, figs 21-32, text-fig. 10. 

Holotype. CNIGR 23/12986, conjoined valves, from the 
Anderken Formation, Sample 2538, Kujandysai section. 

Material. 23 conjoined valves, two ventral and one dorsal valve, 
from Samples 100 (=K-98/1970), 626 Anderkenyn-Akchoku sec- 
tion; Samples 628, 2538 (BC57363), 8257, Kujandysai Section; 
Sample 948 (BC 57314-20), Tesik River. 

Discussion. This species was described and discussed in detail by 
Popov etal. (1999). 



Order ATHYRIDIDA Boucot, Johnson & Staton, 1964 

Suborder ATHYRIDIDINA Boucot. Johnson & Staton. 1964 

Superfamily MERISTELLOIDEA Waagen, 1883 

Family MERISTELLIDAE Waagen, 1883 

Genus KELLERELLA Nikitin & Popov in Nikitin, Popov & 

Holmer, 1996 

TYPE SPECIES. Kellerella ditissima Nikitin & Popov in Nikitin et 
al. (1996), from the Dulankara Regional Stage (Upper Caradoc), 
north Betpak-Dala, Kazakhstan. 

Kellerella misiusi Popov, Nikitin & Sokiran, 1999 

PL 14, figs 58-61 

Holotype. CNIGR 26/1 2986, conjoined valves from the Anderken 
Formation, Sample 2538, Kujandysai section. 

Material. 72 pairs of conjoined valves and one dorsal valve, from 
Samples 100 (=K-98/1970), 626, Anderkenyn-Akchoku section; 
Sample 8214 (BC56773, 57334-9), west side of Aschisu River; 
Samples 628. 2538, Kujandysai Section. 



Table 29 Measurements of ventral valves of Nikolaispira guttula sp. nov.. Sample 948, Tesik river. 





L 


W 


T 


Ld 


Sw 


St 


LAV 


T/L 


Sw/W 


N 


13 


13 


13 


13 


13 


13 


13 


13 


13 


X 


6.5 


5.6 


4.2 


6.0 


2.9 


1.2 


117% 


64% 


52% 


s 


0.72 


0.57 


0.66 


0.70 


0.45 


0.29 


5.2 


5.1 


5.8 


MIN 


5.6 


4.9 


2.8 


5.0 


2.2 


0.5 


110% 


50% 


42% 


MAX 


7.6 


6.7 


5.3 


7.2 


3.6 


1.6 


127% 


70% 


62% 



UPPER ORDOVICIAN BRACHIOPODS FROM KAZAKHSTAN 



77 



cruralium 



median sepum 




Fig. 21 Transverse serial sections of Nikolaispira guttula sp. nov., 1-3, BC 57588; 4-5, BC 57589, Sample 948, Tesik River. Distance in mm is measured 
from the posterior tip of ventral beak. Dorsal valve uppermost. 



DISCUSSION. This species was described and discussed in detail by 
Popov etal. (1999). 



Genus NIKOLAISPIRA Nikitin & Popov in Nikitin, Popov & 
Holmer, 1996 

TYPE SPECIES. Nikolaispira rasilis Nikitin & Popov in Nikitin et 
al., 1996, from the Dulankara Regional Stage (Upper Caradoc), 
north Betpak-Dala, Kazakhstan. 

Nikolaispira guttula sp. nov. PI. 14, figs 47-57; Fig. 21 

ETYMOLOGY. After guttula, Latin - small drop. 

HOLOTYPE. BC 57585, PI. 14, figs 50-53, conjoined valves, from 
the Anderken Formation, Sample 2538, Kujandysai section. 

Material. 38 conjoined valves from Samples 100 (=K-98/1970), 
626, 8221 (BC 56770, 37857-58), Anderkenyn-Akchoku section; 
Samples 628, 2538 (BC 57584, 85), 8257, Kujandysai Section; 
Sample 948 (BC 57588, 89), Tesik River. 

Description. Shell smooth, ventribiconvex, slightly elongate, 
subpentagonal in outline, about 64% as thick as long and 117% as 
long as wide. Anterior commissure parasulcate. Ventral valve profile 
strongly and evenly convex with maximum thickness near the mid- 
valve. Delthyrium small, open, narrow triangular. Beak slightly 
acuminate, erect posteriorly. Shallow ventral sulcus originating 
slightly posterior of the mid-valve, flanked by two low, rounded 
plications and terminating in a narrow, semicircular tongue. Dorsal 
valve gently convex with maximum thickness slightly posterior to 
mid-length. Median fold low and narrow, originating near mid- 
valve. Ventral interior with delicate teeth and short, thin dental plates 
placed closely to the lateral sides of the valve. Dorsal interior with 
small cruralium on a thin, long median septum extending anteriorly 
to mid- valve (Fig. 21). Spiralia laterally directed comprising up to 
three whorls. Jugal processes short, situated near the bese of spiralia. 

DISCUSSION. This species differs from Nikolaispira rasilis Nikitin 
& Popov {in Nikitin, Popov & Holmer 1996), which occurs in the 
Dulankara Stage ( Upper Caradoc to Lower Ashgill ) of north Betpak- 
Dala, Kazakhstan, in having a more elongate outline, like the most 
elongate specimens of the latter species, and smaller number of 
whorls of the spiralia. 



Acknowledgments. We thank Rong Jia-yu (Nanjing) for discussion on 
Chinese material and M.G. Bassett for helpful comments on the manuscript. 
LEP acknowledges support from the Royal Society of London and the 
National Museum of Wales. LRMC acknowledges travel funds from The 
Natural History Museum. 



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1 

Fossil pseudasturid birds (Aves, Pseudasturidae) from the 
London Clay — Novocrania, a new name for the genus 
Neocrania Lee & Brunton, 1986 (Brachiopoda, Craniida), 
preoccupied by Neocrania Davis, 1978 (Insecta, Lepidop- 
tera) — The Creswellian (Pleistocene) human upper limb 
remains from Gough's Cave (Somerset. England) — Gough's 
Cave 1 (Somerset, England): a study of the hand bones — A 
revision of the English Wealden Flora, HI: Czekanowskiales. 
Ginkgoales & allied Coniferales. 2001. Pp. 1-82. £43.40 

The Cenozoic Brachiopod Terebratula: its type species, 
neotype, and other included species. D.E. Lee. C.H.C. Brunton, 
Emma Taddei Ruggiero, Massimo Caldara & Oronzo Simone. 
Gough's Cave 1 (Somerset. England): a study of the pectoral 
girdle and upper limbs. S.E. Churchill. 
Systematic affinity of Acroporella assurbanipali Elliott 
(Dasycladaceae), with notes on the genus Neomeris. Filippo 
Barattolo & Roberta Romano. 

Palynological zonation of Mid-Palaeozoic sequences from the 
Cantabrian Mountains, NW Spain: implications for inter- 
regional and interfacies correlation of the Ludford/Ph'dolf and 
Silurian/Devonian boundaries, and plant dispersal patterns. John 
B. Richardson, Rosa M. Rodriguez, Stuart J. E. Sutherland. 
2001. Pp. 83-162. £43.40 



CONTENTS 



1 Gough's Cave 1 (Somerset, England): a study of the axial skeleton 

S.E. Churchill and T.W. Holliday 
13 Upper Ordovician brachiopods from the Anderken Formation, Kazakhstan: their ecology and 
systematics 

L.E. Popov, L.R.M. Cocks and I.F. Nikitin 




Vol.58, No. 1, June 2002 



ISSN 0968-0462 



Bulletin of 

The Natural History 

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\ tH£ NATURAL 
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VOLUME 58 NUMBER 2 28 NOVEMBER 2002 



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World list abbreviation: Bull. nat. Hist. Mus. Lond. (Geol.) 

ISSN 0968-0462 

The Natural History Museum Geology Series 

Cromwell Road Vol. 58, No. 2, pp. 81-168 

London SW7 5BD Issued 28 November 2002 

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Printed in Great Britain by Henry Ling Ltd, at the Dorset Press, Dorchester, Dorset 



3ull. nat. Hist. Mus. Lond. (Geol.) 58(2): 81-152 Issued 28 November 2002 

The Lower Lias of Robin Hood's Bay, 

Yorkshire, and the work of Leslie Bairstow 

-Mi.' . ..: . J um 

M.K. HOWARTH 

Department of Palaeontology, The Natural History Museum, Cromwell Road, London SW7 5BD i 

3 1 & 

CONTENTS "t^fci.- ^L . ' ■ 'M ' 



ai^»*,^r>7ii^y^--.*^*^-*''- 1 ^ • -— • . i-- 



Introduction 82 

Leslie Bairstow 82 

Biography 82 

Bairstow's unpublished work 84 

Geological maps 84 

Geological structure of Robin Hood's Bay 93 

Stratigraphical succession 93 

Bed numbers 95 

Detailed succession in Robin Hood s Bay 96 

Lithostratigraphy Ill 

Staithes Sandstone Formation 1 ] 1 

Redcar Mudstone Formation 1 1 1 

Exposures in Robin Hood's Bay now 1 14 

Correlation with previous descriptions 1 14 

Bairstow's ammonite collection 1 15 

Systematic description of the ammonites and nautiloids 1 18 

Family Juraphyllitidae 1 18 

Family Lytoceratidae 1 18 

Family Psiloceratidae 1 19 

Family Schlotheimidae 1 19 

Family Arietitidae 119 

SubfamilyArietitinae 1 19 

SubfamilyAgassiceratinae 123 

Subfamily Asteroceratinae 123 

Family Echioceratidae 125 

Family Oxynoticeratidae 129 

Family Cymbitidae 132 

Family Eoderoceratidae 132 

Family Coeloceratidae 136 

Family Phricodoceratidae 137 

Family Polymorphitidae 137 

Family Liparoceratidae 141 

Family Nautilidae 144 

Biostratigraphy 144 

Acknowledgements 150 

References 150 



SYNOPSIS. Rocks of Lower Liassic (Sinemurian and Lower Pliensbachian) age exposed in Robin Hood's Bay. near Whitby, 
north Yorkshire, are described from the mapping, stratigraphical descriptions and ammonite collections made by Mr Leslie 
Bairstow in the years 1927-1970, and preserved in the Palaeontology Department. The Natural History Museum, London. His 
large-scale map of the geology of the foreshore is published on five sheets at a scale of approximately 1 :5000. The stratigraphical 
sequence from bed 4 1 8 at the base up to bed 600.5 at the top of the Lower Pliensbachian is 1 63 .74 m thick, and consists of the 
Redcar Mudstone Formation, for which four members are formally defined - the Calcareous Shale (at the base), Siliceous Shale, 
Pyritous Shale and Ironstone Shale Members - overlain by the lower part of the Staithes Sandstone Formation. The lowest beds 
exposed by the lowest spring tides are Sauzeanum Subzone, Semicostatum Zone, in age; ammonites occur in all subzones, and 
the only uncertain boundary is that between the Masseanum and Valdani Subzones (Ibex Zone), where there are few characteristic 
ammonites. Bairstow's ammonite collection consists of more than 2360 specimens, all from recorded horizons, and is notably rich 
in Promicroceras, Asteroceras, Eparietites and Oxynoticeras from the Obtusum and Oxynotum Zones. Echioceratids, Eoderoceras 
and Apoderoceras from the Oxynotum, Raricostatum and Jamesoni Zones, and Liparoceratids from the Davoei Zone, making it 
a primary source for Sinemurian and Lower Pliensbachian ammonite biostratigraphy. The recently proposed selection of Wine 



i© The Natural History Museum, 2002 



82 



M.K. HOWARTH 



Haven at the south-eastern end of the bay as the Global Stratotype Section and Point (GSSP) for the base of the Pliensbachian Stage 
(ie. the world standard definition), is supported by the sequence of ammonites across the Sinemurian/Pliensbachian boundary. All 
previously figured ammonites from Robin Hood's Bay are listed in a systematic section that includes the evidence on which the 
ammonite identifications in the paper are based, and 56 of the best preserved ammonites are figured. Eparietites bairstowi sp. nov. 
is proposed for an early species of Eparietites and a Sowerby Collection ammonite from the Aplanatum Subzone, Raricostatum 
Zone, in the bay. is designated neotype of Eoderoceras armatum (J. Sowerby). 



INTRODUCTION 



The geology of Robin Hood's Bay (Fig. 1 ) has received the attention 
of many geologists since the 1820s. One such geologist was Leslie 
Bairstow, who decided to make the description of the outcrops of the 
Lower Lias on the foreshore of the bay (Fig. 2) the main scientific 
work of his life. He started serious investigations in 1 928 and worked 
in the bay for the next 50 years. Dr L.F. Spath, a colleague at the 
British Museum (Natural History) (now the Natural History Museum, 
London), identified the many ammonites that he collected and 
brought to the Museum. As long ago as 1956 Spath (1956: 147) 
referred to 'the (still undescri bed) collections made by MrL. Bairstow 
in Robin Hood's Bay', but Bairstow was never able to finish a 
detailed account for publication, and finally he left his work for me 
to complete. That completion has involved more fundamental work 
than mere editing: complete rewriting of the stratigraphical section, 
revision of the maps, preparation of many tables and diagrams not 
envisaged by Bairstow, and revision of the determinations in order to 
produce an up-to-date account of the ammonites and the 
biostratigraphy, were all found to be necessary. The final result is 
eminently worth publishing, if only because it would be very diffi- 
cult to duplicate the ammonite collection, which is the core of the 
paper and the biostratigraphy, at the present time. I had many 




Fig. 1 Map showing the location of Robin Hood's Bay on the north-east 
coast of England, 10 km south-east of Whitby. 



conversations with Bairstow during the period 1956 to 1965,andless 
frequently up to the early 1980s, and quotations from a few of them 
are given here. This paper is not the same as Bairstow would have 
written - his account would have had more local details of the 
outcrops in the bay as they were in the 1920s and 1930s, while the 
paper now presented is more orientated towards correlation by 
ammonites, for which his accurately documented sequence in the 
bay is of major international importance. The comparison that it will 
afford with rocks of the same age and the sequence of zones and 
subzones on the Dorset coast is long overdue. Such comparison is 
too lengthy to be included here - it would involve much collation and 
reidentification of the many separate collections of Dorset ammo- 
nites that now exist, in order to produce a consistent set of 
determinations, and hence biostratigraphy, that could be compared 
with the sequence in Yorkshire. 

The term Lower Lias is used in the title of this paper and elsewhere 
as an exact equivalent of Hettangian + Sinemurian + Lower 
Pliensbachian. This is the sense in which it was widely used and 
understood by palaeontologists when Bairstow worked in the 1920s 
and 1930s. Even in those days a different usage by those geologists 
more interested in lithology and sedimentation led to confusion on 
occasions: the boundary between Lower and Middle Lias was placed 
by them at a position that best marked the change from dominantly 
clayey beds below to dominantly sandy beds above. Such a change in 
lithology occurs at different horizons in different parts of Britain, so 
their usage of Lower and Middle Lias did not have an accurate date 
or age connotation. So much confusion resulted from these disparate 
usages that the terms are rarely used now-a-days. Lower Lias is 
retained here, in the sense given above, in deference to Bairstow and 
the long history of Robin Hood's Bay geology, where it is well 
understood as being the Sinemurian + Lower Pliensbachian, there 
being no Hettangian exposed in the bay. In this usage the Middle Lias 
is exactly equivalent to the Upper Pliensbachian, and the Upper Lias 
to the Toarcian. 



LESLIE BAIRSTOW 

Biography 

Leslie Bairstow (Fig. 3) was born in Halifax, Yorkshire, on 14 August 
1907. After attending Ack worth School (1918-22) and Bootham 
School, York (1922-25), he went to King's College, Cambridge, 
where he obtained a degree in Geology in 1928. He started research 
on the Lower Lias of Robin Hood's Bay in the summer of 1928, 
supported by college scholarships. He had become interested in 
collecting fossils from the Yorkshire Lias during his school and 
undergraduate years, and in getting his ammonites identified he 
came into contact with Dr L.F. Spath at the British Museum (Natural 
History). It was at Spath's suggestion that he decided to undertake 
serious research on the Lower Lias of Robin Hood's Bay, and 
Dr W.D. Lang, also at the Museum, was keen to get another Lower 
Lias section accurately documented for comparison with his own 
work on the Lower Lias of Dorset. He also consulted S.S. Buckman, 
who advised him to record the location of every ammonite he 



LOWER LIAS OF ROBIN HOOD'S BAY 



83 




Fig. 2 The Robin Hood's Bay foreshore at low tide, looking north-west from the top of Ravenscar (above Peak) at the south-eastern end of the bay. Robin 
Hood's Bay town is immediately above the foreshore in the centre-right of the photograph. M.K. Howarth photograph. 28 September 1999. 



collected with sufficient accuracy to enable the sequences of ammo- 
nite 'hemerae' to be compared at the north-western and south-eastern 
ends of Robin Hood's Bay. Initially the project was intended to be a 
thesis for a higher degree at Cambridge University, but the detailed 
mapping, bed-by-bed description and collecting during 1928-1930 
were submitted as a dissertation in support of a fellowship applica- 
tion at King's College in late 1930. This was not successful, and 
Bairstow was preparing for a second application in 1931, when the 
offer of a permanent post at The Natural History Museum in South 
Kensington (then the British Museum (Natural History)), with the 
opportunity to continue work indefinitely on the Lower Lias of 
Robin Hood's Bay, appealed to him more than a fellowship at 
Cambridge of six years duration. In fact, during his early years at the 
Museum he was elected to a three-year visiting Fellowship at King's 
College in 1 932-35 . He started at the Museum in October 1 93 1 , and 
although initially put in charge of fossil echinoderms and later 
Coleoidea (including belemnites), he was able to continue work on 
Robin Hood's Bay until his retirement in June 1 965. He continued to 
work at the Museum until 1985, when he moved to Todmorden, 
Yorkshire, where he died on 10 August 1995. 

The meticulous attention to detail that Bairstow lavished on the 
description, collecting and mapping of the Lower Lias of Robin 
Hood's Bay, made it unlikely that he would produce a final descrip- 
tion with which he would be satisfied. It is fortunate, therefore, that 
for the Cambridge fellowship dissertation of 1930 he produced a 
finished manuscript version of the map and a stratigraphical descrip- 
tion of the whole succession, which are the basis of the present paper. 
His most useful collecting occurred during the period 1928-34, and 
work during the following 40 years did not greatly enhance that 
original burst of activity. His ammonite collection of about 2,360 



specimens is housed in The Natural History Museum, and is a prime 
record of the sequence of ammonite faunas for the upper two-thirds 
of the Lower Lias. Such a collection would be difficult to repeat 
today, because so many of the accessible ammonites have been 
removed from the Bay. Bairstow conducted field parties to the Bay 
for the 18th International Geological Congress in 1948 and the 
William Smith Jurassic Symposium in 1 969, and gave brief summa- 
ries of the zonal sequence and his bed numbering in the guide books 
for those meetings (Bairstow, 1948, 1969). Apart from a summary in 
a guide to the fossils of the Scarborough district (Bairstow, 1953), he 
left no other published account. 

The geological map of Robin Hood's Bay, the description of the 
stratigraphy and basic biostratigraphy formed the first half of his 
dissertation for the fellowship at King's College, Cambridge, in 
1930. The second half of that dissertation consisted of a description 
of the Lower Lias belemnites of Robin Hood's Bay, and included an 
assessment of the 19 specific names proposed by Simpson (1855: 
22-3 1 ; 1 884: 47-54) and partly revised by Phillips ( 1 863-1 909), for 
belemnites from the bay. This part of his work was also destined 
never to be published, but it did give Bairstow an interest in belemnites 
and related groups, which led to him being put in charge of fossil 
Coleoidea at the Natural History Museum. In fact, he did much 
valuable investigation into the early generic nomenclature of fossil 
Coleoidea, especially the non-belemnite groups, and his detailed 
notes were passed on first to Dr J. A. Jeletzky, then to later Treatise 
authors, for incorporation in the Coleoid volume of the Treatise on 
Invertebrate Paleontology (not yet published). In these and other 
matters, especially the geology of Robin Hood's Bay and general 
identification of specimens sent to the Museum, colleagues found 
him helpful and were always enlightened by his views. He had a long 



84 



M.K. HOWARTH 




Fig. 3 Leslie Bairstow, aged 46; taken from a group photograph of the 
Palaeontology Department, the Natural History Museum, 1954. 



association with the Palaeontographical Society, first as Treasurer 
for the years 1948 to 1955, then as Vice President from 1966 until 
1969, and he was a Trustee of that Society for several years during 
the same period. [Other biographies on Bairstow can be found in the 
obituary notices published by the Geological Society (Howarth, 
1996) and King's College. Cambridge (Annual Report, October 
1996:27-29).] 



Bairstow 's unpublished work 

The originals of the unfinished and unpublished manuscripts left by 
Bairstow will be deposited in the Earth Sciences Library of The 
Natural History Museum, London. They can be divided into three 
main parts, which will now be described in sequence: 

1. The geological map of Robin Hood's Bay. 

2. The stratigraphical description of the Lower Lias. 

3. His collection of ammonites and other fossils, and his list of the 
determinations of the ammonites. 



GEOLOGICAL MAPS 

Bairstow drew his geological map of Robin Hood's Bay in 1928-30 
and the top copy formed part of his fellowship submission, now 
preserved in the archives of King's College, Cambridge. The map 
was drawn at a scale of 1:2500, and consists of eight sheets, each 
measuring 320 x 343 mm. Bairstow took (and variously modified) a 



few geographical lines from the 1:2500 sheets of the Ordnance 
Survey: these were mainly the top and bottom of the cliffs, some 
paths and field boundaries at the top of the cliffs, a few roads and 
prominent buildings, and the line of low tide mark. The latter is the 
low tide mark of ordinary tides on Ordnance Survey maps (ie. 
approximately half way between low water mark of spring and neap 
tides), but Bairstow modified the line to be that of low water mark of 
spring tides, when the maximum amount of rock is exposed on the 
foreshore. On some low lying areas low water of spring tides exposes 
much larger areas of rock than ordinary or neap tides, eg. the area 
occupied by beds 505-530 north of Robin Hood's Bay town. 

Before submission to King's College. Bairstow made machine 
copies of his map, which he kept throughout his working life and 
they form the basis of the maps reproduced here. These copies differ 
from the original maps only in Bairstow's addition of 'datum lines' 
for locating the ammonites he collected (see the account of his 
ammonite collection, p. 1 17 below). I considered the possibility of 
reproducing Bairstow's original maps in the King's College archives, 
but the small size of the lettering of the bed numbers, the colour, 
thinness and lack of sufficient boldness of some of the lines, and 
logistical difficulties of reproducing maps that are larger than A3 (in 
one direction), were all against direct reproduction of the originals. 
After consideration of scale and legibility at the final printed size, it 
was decided to copy the maps at a different scale, and to divide them 
into the five sheets (see Fig. 4) that give a better division of the 
outcrops in Robin Hood's Bay than the original eight maps. Tracing 
was done with great care, so that the geological lines on the resulting 
maps reproduced here as Figs 5, 6. 8. 11 and 15 are as close as 
possible to the lines originally drawn by Bairstow. No alterations 
were made, and in those parts that were checked, such as the seaward 
edge of outcrops opposite the mouth of Mill Beck, south of Stoupe 
Beck, and in Wine Haven, the maps appear to be still accurate, 70 
years after they were drawn. 

Bairstow did not. however, include the lowest and highest beds on 
his map: the lowest bed he mapped was the seaward edge of bed 422, 
Low Balk, though he collected ammonites from lower beds down to 
bed 42 1 . 1 . The outcrops of beds 4 1 8-42 1 , seaward of Low Balk, were 
added to the map of Fig. 1 1 from observations made at the lowest 
spring tide of the year on 9 September 1991. when they were easily 
accessible. Similarly, at the top of the succession, the highest bed 
mapped by Bairstow was bed 590.3, on the south side of Bulmer Steel 
Hole, even though this is 250 m south of Castle Chamber, which is 
usually taken as marking the northern boundary of the bay. The higher 
beds northwards past Castle Chamber and up to the boundary between 
the Lower and Upper Pliensbachian ( ie. the Lower/Middle Lias boun- 
dary ) were also added to the map of Fig. 5 in September 1 99 1 . Thus, the 
mapping was completed between the lowest bed exposed by the lowest 
spring tides and the upper boundary of the Lower Pliensbachian. The 
five large-scale maps are printed here at a scale of 1:5315. 

Map 1 (Fig. 5) is the northernmost map and starts from the highest 
beds outside the bay to the north-west at the junction with the Upper 
Pliensbachian. The main geographical feature is the cave of Castle 
Chamber, where the hard shelly sandstones of beds 599 and 601.1 
form the floor and roof of the cave. The outcrop is fairly narrow along 
the whole of this east facing part of the bay, and is subject to 
aggressive wave action that results in relatively clean rock surfaces 
and good exposures. 

Map 2 (Fig. 6) covers the whole sweep of the scars north and south 
of Robin Hood's Bay town, from the ironstone shales at the top of the 
map, down through the softer, pyritous shales opposite Dungeon 
Hole and Ground Wyke (Fig. 7), to the hard siliceous shales that form 
prominent scars opposite Robin Hood's Bay town. These are Landing 



LOWER LIAS OF ROBIN HOOD'S BAY 



85 



-CASTLE CHAMBER 



BULMER STEEL 
HOLE 



NESS POINT 
I or 
NORTH CHEEK 

I 

I 
J 




07- 



ZONE 


SUBZONE 




Margaritatus 


Stokesi 


Sto 


STAITHES 

SANDSTONE 
FORMATION 


Davoei 


Figulinum 


Fig 


Capricornus 


Cap 


Maculatum 


Mac 


Ironstone 

Shale 

Member 


Z 
O 
H 

< 

s 

2 
o 
n, 

w 

z 
o 

H 

§ 

< 
u 

Q 


Ibex 


Luridum 


Lur 


Valdani 


Val 


Masseanum 


Mas 


Jamesoni 


Jamesoni 


Jam 


Brevispina 


Bre 


Polymorphus 


Pol 


Taylori 


Tay 


Pyritous 

Shale 

Member 


Raricostatum 


Aplanatum 


Apl 


Siliceous 

Shale 

Member 


Macdonnelli 


Med 


Raricostatoides 


Rar 


Densinodulum 


Den 


Oxynotum 


Oxynotum 


Oxy 


Simpsoni 


Sim 


Obtusum 


Denotatus 


Dnt 


Stellare 


Ste 


Obtusum 


Obt 


Calcareous 

Shale 

Member 


Turneri 


Birchi 


Bir 


Brooki 


Bro 


Semicostatum 


Sauzeanum 


Sau 


Scipionianum 


Not exposed 


Reynesi 



1 Km 

_J 



06- 



05- 



04- 



03- 



Fig. 4 Summary geological map of the foreshore in Robin Hood's Bay, showing the main geographical features. The geological divisions shown are the 
subzones, and the areas covered by the five main maps of Figs 5.6,8, II and 1 5 are indicated by the rectangles of dashes. The cliff is indicated by 
vertical lines showing approximately the steepest direction of the face. The table containing the key to the subzones is a summary of Table 1. 



86 



M.K. HOWARTH 



MAP 1 


i 
N 


\ Up P er ( 
V \ Pliensbachian V\ 8 s 5 

^C?c& V s \\VV ^ 


1 

96 

1 / V 










\ a > 


\\i 1 ^ \ 


CASTLE 








\ \ 


v Ik J 


Chamber 








i 


1 


300 m \ / i 

I l \ I 


/// ^ 


•5*" 

^ 












\ 


J/ /$/ 


BULMER 








r'i 


STEEL 
HOLE 






il 


V / / ^y 

u r//A 


,589 






1 1 


J III & ( 


~\^^ 






'•( 


/ ^ %b / 


^2-5&5 






i ! 1 




^5» 

5SY NESS POINT 
\ or 






\ !( y 


y^t^^/ 


L.S19 

NORTH 






i'j // 




CHEEK 


-06 




-y\%£yyyfT M 


;^?-?\ "576.5 -.9 

^ / 576.4 

^/-575 

-573 


.578.4 

-NESS RUCK 

06- 




't*^ 


A /-W-^jS^j? 21 - 562 

/ ^-^ ^-^y^y 561 

^/^y^^V^^st-- 56 ° ' 


96 

1 





Fig. 5 Map 1 . the northernmost part of Robin Hood's Bay, showing the outcrops up to the north-west corner of the bay round to the junction with the 
Upper Pliensbachian (Middle Lias). 



LOWER LIAS OF ROBIN HOOD'S BAY 



87 




Fig. 6 Map 2, showing outcrops down to the bottom of the Lower Pliensbachian and into the top of the Upper Sinemurian in the rock scars southwards 
past Robin Hood's Bay town. 



M.K. HOWARTH 




it sraHf 



'mz 



Fig. 7 The foreshore at low tide north of Robin Hood's Bay town. L. Bairstow photograph. 1952. taken from the top of the cliff near the northern edge of 
Fig. 6. The rock outcrops can be seen to have been relatively clean and free of algae and beach deposits at this time. 



Scar' (bed 496) and East Scar (bed 494) opposite the town, and 
Cowling Scar (bed 474, Double Band) farther out to sea near the 
bottom edge of the map. The relatively soft beds of the Pyritous 
Shales around Ground Wyke are the wettest and lowest (relative to 
sea level) exposed part of the foreshore in the bay, where there is now 
little or poor rock exposure owing the seaweed, barnacle and mussel 
bed cover. 

Map 3 (Fig. 8) shows an interesting entity of outcrops around the 
mouth of Mill Beck, the cliffs that make up 'The Nab' . and the major 
rock scars on the foreshore to the east. There are three prominent 
scars here - Low Scar, Middle Scar and High Scar, being the hard, 
calcified silty shales of beds 447, 449 and 455 respectively. Also 
notable on this map are Tinkler's Stone and Strickland's Dumps, 
north of the mouth of Stoupe Beck, and both are named on the larger 
scale Ordnance Survey maps. Tinkler's Stone lies on bed 462 and is 
a boulder of very hard grey-brown massive limestone, of undeter- 
mined origin, but not derived from the Lower Lias. Strickland's 
Dumps are small, but relatively deep, excavated pools in the dip 
slope of beds 455.1 and 455.2. The area around Bay Mill and The 
Nab is shown on a larger scale in Fig. 9. High tides penetrate well into 
the mouth of Mill Beck between The Nab and the road on the south 
side of the beck, and sometimes large masses of dead algae partly 
block or divert the outflow of Mill Beck. However, when the mouth 



"Scar' is a term frequently used in descriptions of Yorkshire coast geology for rock 
outcrops on the foreshore, which are usually below (but may sometimes be above) the 
level reached by normal high tides. In Robin Hood's Bay several scars are given formal 
names on some larger scale maps of the Ordnance Survey, and a few others are newly 
named in this paper. Such scars are formed by, or at least topped by, single beds of hard 
rock, and all are intertidal in Robin Hood's Bay. 



and bed of Mill Beck are clear of algae, the underlying beds can be 
seen from bed 475 at the mouth of the beck, past Bay Mill and up to 
bed 494 in front of the weir. They were mapped by Bairstow as 
shown on Fig. 9 in an amount of detail that is too great to be shown 
clearly on the main map. The face of The Nab. with some individual 
beds identified, is shown in Fig. 10. 

Map4(Fig. 1 1) features the foreshore north-east of Peter White Cliff 
where there are exposures down to the lowest beds in Robin Hood's 
Bay (Fig. 12). At low tides the scars of Low Balk (bed 422; see Fig. 
13) and the slightly less conspicuous Pseudo Low Balk (bed 424.2) 
are prominent rock platforms, both of which can normally be reached 
only by wading through the shallow channels between beds 424 and 
425 and along the middle part of bed 422.2, which never dry out, 
even at the lowest spring tides. Bed 447 forms a long scarp face 
across the whole width of the map in front of Peter White Cliff (Fig. 
14). On the Ordnance Survey maps, the name High Scar is used for 
two different beds: one for this bed 447 in front of Peter White Cliff, 
the other for bed 455 east of Mill Beck. So bed 447 forms both Low 
Scar east of Mill Beck and the Ordnance Survey's 'High scar' at 
Peter White Cliff. In view of the possible confusion, the latter use of 
'High Scar' is not perpetuated here. 

Map 5 (Fig. 15) reaches the south-east corner of the bay, where the 
Lower Lias succession is truncated by the Peak Fault complex. The 
Main Peak Fault has a downthrow to the east and a large lateral 
movement, resulting in Upper Pliensbachian and Toarcian beds on 
the east abutting the top beds of the Sinemurian and bottom beds of 
the Lower Pliensbachian on the west. There is a narrow zone of 
severely shattered beds immediately west of the fault, and minor 
faults and cracks for some distance farther west. The highest bed 
exposed on the rock platforms below the cliffs is bed 501.1, the 



LOWER LIAS OF ROBIN HOOD'S BAY 



89 



MAP 3 



300 m 




04- 



Fig. 8 Map 3, the middle map of the bay, showing the prominent scars on the foreshore east of The Nab and the mouth of Mill Beck, down to the mouth of 
Stoupe Beck at the bottom of the map. 



90 



M.K. HOWARTH 




Fig. 9 Details of the outcrops of beds 475^4-94 in the bed of Mill Beck from the mouth of the beck up to the weir west of Bay Mill. 




Fig. 10 The seaward face of The Nab immediately north of the mouth of Mill Beck. M.K. Howarth photograph. 1 1 September 1991. The beds here are the 
middle part of the Siliceous Shale Member and belong to the Oxynotum to basal Raricostatoides Subzones; the main beds are identified on the right hand 
side of the photograph. 



LOWER LIAS OF ROBIN HOOD'S BAY 



91 



MAP 4 




o 



300 m 

I 



96 



Fig. 11 Map 4, showing outcrops down to the lowest beds exposed in the bay north-east of Peter White Cliff. 



92 



M.K. HOWARTH 




2.6 F. Sh* 16 . 4*522..?. 10- 10-38 10+0 ra.irt. tooo tt S.v/. 



Fig. 12 Oblique aerial view of the foreshore in front of Peter White Cliff at low tide. Air Ministry photograph. 10 October 1938. formerly Crown 

Copyright. The line of Low Balk (bed 422.2) is the lowest bed visible farthest from the cliff (see Fig. 13). and the long outcrop of the prominent bed 447 
can be seen just below the cliff (see Fig. 14). 




Fig. 13 Low Balk (bed 422.2) at low water of spring tide, almost the farthest accessible point north-east of Peter White Cliff. M.K. Howarth photograph. 
10 September 1991. 



LOWER LIAS OF ROBIN HOOD'S BAY 



93 




Fig. 14 The prominent hard calcified shale of bed 447 in front of Peter White cliff. M.K. Howarth photograph. 10 September 1991. 



lowest bed of the Lower Pliensbachian. The most prominent bed on 
the map is bed 474, Double Band, which forms Billet Scar (see Fig. 
16). The narrow excavation through beds 476^186 known as The 
Dock, was originally made for fishing and smuggling purposes. 



GEOLOGICAL STRUCTURE OF ROBIN 
HOOD'S BAY 

The pattern of the outcrops on the foreshore of the bay as seen in Fig. 
4 is determined by the structure of the rocks (Fig. 1 7 ) . That structure 
was first alluded to by Tate & Blake (1876: 27, 196) who described 
Robin Hood's Bay as 'a complete inlier ... in the form of a mound, 
dipping in all directions from the centre . . . the centre of elevation 
beneath the sea, nearly opposite the centre of the bay'. In the 
Geological Survey memoir, Fox-Strangways & Barrow (1915: 3, 
115) referred to the Lias as 'curving over in a gentle arch or 
anticline'. Versey (1939: pi. 15) plotted the contours of the base of 
the Grey Limestone (=Scarborough Formation; Lower Bajocian) 
over a wide area and showed that around the southern and western 
sides of Robin Hood's Bay they formed the outer part of a north-west 
to south-east elongated dome . According to Versey (1939) the dome 
was produced by tectonic movements probably in the late Pliocene. 
Kent (1974: 25, 26) and de Boer (1974: 281) accepted the date of 
formation of the dome as later than the mid-Tertiary Alpine move- 
ments and probably Pliocene. 
The central part of the Robin Hood's Bay dome can be defined by 



the outcrops of the Lower Lias on the foreshore of the bay. Fig. 17 
shows contours on the outcrop at 10 m bed-thickness intervals, with 
the m contour starting at the top of bed 422.2, Low Balk. Because 
the outcrop on the foreshore is essentially flat, the contours approxi- 
mate closely to strike lines, and they form the pattern of a dome, with 
a NW to SE axis of elongation in approximately the position shown 
on the figure. The dip of the beds is at right angles to each contour 
line away from the centre of the dome. In the northern part of the bay 
the beds dip NW. and from the 50 m to the 150 m contours the 
average distance between adjacent contours is 128 m; this gives an 
average dip of 4.5° for the beds. Between the 20 m and 40 m contours 
the beds dip to the west, while the lowest beds between the m and 
20 m contours dip between west and south at an average of 3.2°. In 
the south-east corner of the bay near the Peak Fault the beds curve 
round to dip south-easterly. The Peak Fault throws down on its 
eastern side and has a lateral movement of several kilometres. Its 
northern continuation across Robin Hood's Bay, passing close to the 
shore at the northern end of the bay, is shown on Fig. 1 7 in accord- 
ance with data on the latest map of the British Geological Survey that 
includes off-shore geology (British Geological Survey, 1995, Tyne 
Tees, sheet 54°N-02°W, 1:250,000, solid geology). 



STRATIGRAPHICAL SUCCESSION 

Bairstow drew up a description of the succession in Robin Hood's 
Bay in 1928-30 and the detailed sequence of beds formed an 



94 



M.K. HOWARTH 




LOWER LIAS OF ROBIN HOOD'S BAY 



95 




Fig. 16 The foreshore at low tide in Wine Haven looking eastwards towards Peak and the bottom of the cliff below Ravenscar. L. Bairstow photograph, 
1929 or 1930. This view looks along the prominent outcrop of Billet Scar (bed 474) in the middle of the photograph, and 'The Dock' crosses the beds 
obliquely to the right. 



important part of his fellowship submission to King' College. He 
kept carbon copies of the 85 typed pages of the sequence, and during 
subsequent years up to 1 975 he made alterations, additions and notes 
on the originals until the final size of the manuscript was about 230 
pages. Many alterations were made to the bed numbering, especially 
at the top, and to the bed thicknesses and details of the lithology, none 
of which were fully finalized at the time of his fellowship submis- 
sion. This manuscript is the basis for the much edited version of the 
stratigraphical succession given below, where as much of the 
lithological description as necessary has been retained to describe 
and identify individual beds in the sequence. Bairstow measured bed 
thicknesses on both the foreshore scars and in the cliffs in order to 
arrive at figures he considered accurate, and his measurements in 
feet and inches have been converted to metric units for this paper. 

The lithostratigraphical divisions given in the succession below 
are not those of Bairstow. They are based on more recent work 
described below in a separate section. Similarly, the zone and 
subzone divisions given in the succession are based on revisions of 
the identifications of all Bairstow's ammonites, also as described in 
a separate section. Table 1 shows a detailed correlation between the 



zones and subzones, the bed numbers and the lithostratigraphical 
divisions. 

Bed numbers 

Bairstow started his detailed description of the beds in 1928 by 
giving the bed number 500 to the nodules that form the northern 
boundary of The Landing at Bay Town, and worked up and down the 
succession from that level. That bed number was selected because he 
did not know what his lowest and highest numbers would be, and 
also to 'prevent confusion with the numbers [1-132] given by Lang 
to Lower Lias beds of similar age on the Dorset coast' . After several 
changes to his various schemes, especially in the top part of the 
succession, he finalized his numbering with bed 418 as the horizon 
exposed at the lowest level reached by spring tides in the bay, and bed 
601 as the highest he described in the Staithes Sandstone Formation 
just beyond the northern end of the bay. In various places he 
subdivided individual beds by giving numbers after a decimal point 
(eg. beds 485. 1 , 485 .2, 485.3 ), and a few beds were subdivided to two 
places of decimals (eg. beds 464.31, 464.32, 464.33). In bed 590, 



96 



M.K. HOWARTH 



STAGE 


ZONE 


SUBZONE 


m 


BED NO 




U. Pliensbachian 


Margaritatus 


Stokesi (part) 


600.6-601.2 


STAITHES 
SANDSTONE 
FORMATION 


Lower 
Pliensbachian 


Davoei 

32.63 m 


Figulinum 


9.70 


596.2-600.5 


Capricornus 


3.04 


591-596.1 


Maculatum 


19.89 


581-590.7 


Ironstone 

Shale 
Member 
62.73 m 


a 

z 
o 

H 
< 

O 

W 

z 
o 

E-i 

Q 

D 

§ 

< 
U 

1 


Ibex 
20.39 m 


Luridum 


7.24 


578.1-580 


Valdani 


7.66 


571-577 


Masseanum 


5.49 


560.3-570 


Jamesoni 
44.46 m 


Jamesoni 


5.66 


550-560.3 


Brevispina 


3.72 


544.6-549 


Polymorphus 


7.05 


538-544.5 


Taylori 


28.03 


527-537 


501.1-526.7 


Pyritous Shale 
Member, 26.18 m 


Upper 
Sinemurian 


Raricostatum 
17.26 m 


Aplanatum 


5.57 


497-500 


496 


Siliceous 

Shale 
Member 
38.74 m 


Macdonnelli 


4.48 


494-495.7 


Raricostatoides 


6.21 


488-493.5 


Densinodulum 


1.00 


486.3-487 


Oxynotum 
14.91 m 


Oxynotum 


9.19 


472.1-486.2 


Simpsoni 


5.72 


463-471 


Obtusum 
12.45 m 


Denotatus 


3.37 


455.2-462 


Stellare 


7.37 


447-455.1 


Obtusum 


1.71 


446.31-446.5 


Calcareous 

Shale 

Member 

23.35 m exposed 


Lower 
Sinemurian 

(part) 


Turneri 

7.75 m 


Birchi 


5.89 


433.3-446.2 


Brooki 


1.86 


429.7-433.2 


Semicostatum 
13.89 m 


Sauzeanum 


13.89 


418-429.64 


Scipionianum 


Not exposed 


Reynesi 



Table 1 Summary of the bed numbers used in Robin Hood's Bay, and their grouping into zones and subzones (including thicknesses), and members and 
formations, showing the detailed correlation between biostratigraphical and lithostratigraphical divisions. 



however, he used three places of decimals (eg. bed 590.433) and in 
bed 598 he used four places of decimals (eg. bed 598.4322). Three 
and four places of decimals are considered here to be too cumber- 
some to be acceptable, so they have all been replaced in this 
description with the minimum amount of renumbering necessary to 
achieve single and double decimal numbering in beds 590 and 598. 
Unfortunately, it was not possible to replace all the double decimal 
numbering in the succession, because there are more than 9 divisions 
in beds 429, 495, 544 and 590, and to replace them would have 
involved renumbering the whole succession. This was not practica- 
ble in view of the large number of entries of the original bed numbers 
on specimen labels, index cards and original manuscripts and maps. 
It should be noted, however, that the subdivisions that Bairstow used 
for bed 600 are not in a decimal system like those in all the lower beds 
- subdivisions of bed 600 use the 13 suffix numbers 1-13 after the 
decimal point; as these are at the top of the succession extending out 
of Robin Hood's Bay to the north, they are retained here without 
alteration. 



DETAILED SUCCESSION IN ROBIN HOOD'S 
BAY 

In the following detailed succession records of all the ammonites and 
nautiloids in Bairstow's collection are included for each bed; the first 
number in brackets following each species is the total number of 
specimens, and is followed by their registration numbers, then by a 
reference to any specimens figured here; in a few cases the number 
of registration numbers quoted is less than the total number of 
specimens recorded, because specimens were lost, destroyed, poorly 
preserved, uncollectable (but observed by Bairstow), or too numer- 
ous to be worth registering all of them. The thickness of each bed is 
given in the right hand column in metres (m). 

Specimen register numbers are identified here and in the remain- 
der of the paper by the following prefixes: C. and CA - The Natural 
History Museum, London; GSM - British Geological Survey (Geo- 
logical Survey Museum), Keyworth, Nottinghamshire; OUM - 
Oxford University Museum; SM - Sedgwick Museum, Cambridge; 
WM - Whitby Museum, Yorkshire. 



LOWER LIAS OF ROBIN HOOD'S BAY 



97 




Fig. 17 Map showing contours at 10 m bed thickness intervals in the Lower Lias on the foreshore of Robin Hood's Bay, from which the elongated dome 
geological structure can be deduced. The only geographical features shown are the line of the base of the cliff and the low tide mark. See text for further 
explanation. 



STAITHES SANDSTONE FORMATION (PART) 



Bed no 


601.2 


601.1 


600.13 


600.12 


600.11 


600.10 


600.9 


600.8 


600.7 


600.6 



Zone of Amaltheus margahtatus 
Subzone of Amaltheus stokesi (lower part) 

m 

Sandstone, soft, micaceous, laminated 0.66 

Sandstone, hard, shelly in parts; many Gryphaea sp.; forms the roof of Castle Chamber at north-west end of Robin 

Hood's Bay 0.81 

Sandstone, soft, silty 0.33 

Shale, silty 0.38 

Limestone nodules 0.10 

Shale, silty 0.08 

Shale, sandy 0.38 

Shale, silty, with horizon of limestone nodules near top 0.53 

Amaltheus stokesi (J. Sowerby) 0.3 m above base ( 1 ; CA 4605) 

Shale, sandy, laminated; forms the most conspicuous positive feature between floor and roof of Castle Chamber 0.25 

Flat limestone nodules 0.13 

Amaltheus stokesi (J. Sowerby) (1; MKH Coll, lost). 



98 M.K. HOWARTH 

Zone of Prodactylioceras davoei 
Subzone of Aegoceras (Oistoceras) figulinum 

600.5 Shale, silty, pale grey 0.36 

600.4 Limestone nodules (= bed v of Howarth, 1955: 155) 0.06 

Aegoceras (Oistoceras) figulinum (Simpson) (1 1; SM J35968, SM J44776-85). 

600.3 Shale, silty 0.91 

600.2 Limestone, sideritic, forming a continuous bed (= bed iii of Howarth, 1955: 155) 0.18 

Aegoceras (Oistoceras) angulatum (Quenstedt) (9; SM J44790, CA 4595-4602). 

600.1 Shale, silty 0.15 

599 Sandstone, flaggy, with ripple marks and oyster bands; the top surface forms the floor of Castle Chamber (= bed i of 

Howarth, 1955: 155) 1.42 

Aegoceras (Oistoceras) angulatum (Quenstedt) (1; SM J44789) 

598.35 Shale, silty, laminated 0.33 

598.34 Limestone nodules 0.08 

598.33 Shale, silty, laminated 1.00 

598.32 Limestone, grey, weathering yellow; forms continuous tabular bed 0.10 

598.31 Shale, silty, soft, but harder near top 2.79 

598.2 Shale, silty, hard; forms a conspicuous feature in the cliff and on the scar 0.15 

598.1 Shale, silty, soft, darker than the two beds above; occasional sideritic mudstone nodules in middle and upper part 0.67 

Aegoceras (Oistoceras) sinuosiforme Spath (5; C.38930, C.39556, C.39579, CA 4594; PI. 7, fig. 14). 

597 Limestone nodules 0.13 

Aegoceras (Oistoceras) sinuosiforme Spath (42; C.39398, C.39418-36 [C.39421/22 and C. 39429/30 are single 
specimens], C.39499-39502, C.39505-07, C.39561, C.39568-77, CA 4588-93). 

596.3 Shale, silty, with 0.09 m thick limestone nodules, especially in a band 0.3m below the top 1.22 

Aegoceras (Oistoceras) sinuosiforme Spath (28; C.39396-97, C.39399-39417, C.39437-38, C.39504, C.39549, 

C.39559, CA 4586-87), Liparoceras (L.) divaricosta (Trueman) (1; C.39455; PI. 8, fig. 1). 

596.2 Hard, calcified shale, with occasional sideritic mudstone nodules 0.15 

Aegocras (Oistoceras) sinuosiforme Spath (18; C.39503, C.39508-10. C.39560, C.39562-67, C.39578. CA 4580-85). 

Subzone of Aegoceras (A.) capricornus 

596.1 Shale, soft; occasional calcareous mudstone nodules 0.85 

595.2 Shale, silty, lower half soft, upper half harder; some lenses of oysters 0.56 

595.1 Scattered calcareous mudstone nodules in hard silty shale 0.10 

594 Shale, silty, with a few calcareous mudstone nodules 0.61 

Aegoceras (A.) lataecosta (J. de C. Sowerby) (1, lost), Aegoceras (A.) sp. indet. (10; CA 4579). 
593 Sideritic mudstone nodules; conspicuous on the scars at Bulmer Steel Hole 0.13 

Aegoceras (A.) artigyrus (Brown) (4; CA 4604). 

592 Shale, silty, laminated with some hard bands; a few sideritic mudstone nodules 0.61 

591 The Oyster Bed. Hard, silty shale, with flat sideritic mudstone nodules; many oysters and other bivalves 0.18 

Aegoceras (A.) lataecosta (J. de C. Sowerby) (1; C. 39395). 

REDCAR MUDSTONE FORMATION 
IRONSTONE SHALE MEMBER 

Subzone of Aegoceras (A.) maculatum 

590.7 Shale, silty 1.42 

590.66 Spherical sideritic mudstone nodules 0.10 

590.65 Shale, silty, with scattered sideritic mudstone nodules 0.23 

590.64 Sideritic mudstone nodules 0.08 

590.63 Shale, silty, with scattered sideritic mudstone nodules near the base 0.20 

Aegoceras (A.) maculatum (Young & Bird) (8; C.38881. C.38886-90, C.38893, CA 4578), Aegoceras (A.) maculatum 

(Young & Bird) var. leckenbyi Spath ( 1 ; C.38880). 

590.62 Shale, silty 0.41 

590.61 Shale, silty, with many sideritic mudstone nodules 0.46 

Aegoceras (A.) maculatum (Young & Bird) (8; C.38882-85, C.38891-92, C.38894, C.38896; PL 7, fig. 13). 

590.5 Shale, silty 1.30 

590.43 Shale, silty, with a few sideritic mudstone nodules near the top 0.23 

590.42 Calcareous mudstone nodules 0.08 

590.41 Shale, silty, with a few sideritic mudstone nodules near the base 0.20 

590.3 Shale, silty, with occasional very large (up to 1 m diameter) sideritic mudstone nodules 0.5-1.5 m below the top; seen 

on the scar on the south side of Bulmer Steel Hole 3.58 



LOWER LIAS OF ROBIN HOOD'S BAY 99 

590.2 Hard silty shale, with many small sideritic mudstone nodules 0.41 

590.1 Shale, silty, with occasional sideritic mudstone nodules 0.51 

Aegoceras (A.) maculatum (Young & Bird) (7; C.38895. CA 4573-77), lAndwgynoceras sp. indet. (1; CA 4603). 

589 Large sideritic mudstone nodules; a strongly marked band crossing the shore south of Bulmer Steel Hole 0.15 

588 Shale, silty; one very large sideritic mudstone nodule 1 m below top 4.95 

Aegoceras (A.) maculatum (Young & Bird) (5; C.39136, C. 38878-79, CA 4571-72). Androgynoceras heterogenes (Young 

& Bird) (1; C.39136), Liparoceras (L.) sp. indet. (2; CA 4562-63). 

587 Flat septarian sideritic mudstone nodules, some with domed upper surfaces 0.13 

586 Shale, silty 2.29 

585 Flat septarian sideritic mudstone nodules, some with domed upper surfaces, and sometimes forming a continuous bed .... 0.13 

Aegoceras (A.) maculatum (Young & Bird) (1; C.38877). 
584 Shale, silty 1.45 

ILytoceras sp. indet. ( 1 ; CA 2795), Aegoceras (A.) maculatum (Young & Bird) (2; C.38876, CA 4570). 

583.2 Shale, with scattered sideritic and calcareous mudstone nodules 0.08 

Aegoceras (A.) maculatum (Young & Bird) (2; C. 38871-72), Aegoceras (A.) maculatum (Young & Bird) var. atavum 

Spath (2; C.38873-74: PI. 7, fig. 8), Androgynoceras heterogenes (Young & Bird) (1; C.38875). 

583.1 Continuous bed of sideritic mudstone 0.08 

582 Shale, silty, with small scattered spherical sideritic mudstone nodules 0.25 m below top 1.27 

Liparoceras (L.) cf. naptonense Spath (1; C. 39140), lAegoceras s.l. sp. indet. (1; not registered). 
581 Flat sideritic mudstone nodules 0.15 

Aegoceras (A.) maculatum (Young & Bird) (2; C.41307, CA 4569; PI. 7, fig. 12). 

Zone of Tragophy 'Hoc eras ibex 
Subzone of Aegoceras (Beaniceras) luridum 

Shale 1.30 

Aegoceras (Beaniceras) cf. luridum (Simpson) (1; CA 4568), Liparoceras (L.) cf. naptonense Spath (1; C. 39141). 

Flat sideritic mudstone nodules 0.15 

Liparoceras (L. ) sp. indet. ( 1 ; lost). 

Shale 3.40 

Aegoceras (Beaniceras) luridum (Simpson) (6; CA 4565-67), Liparoceras (L.) sp. indet. (1; CA 4561), Lytoceras 

fimbriatum (J. Sowerby) ( 1 ; lost); many large Inoceramus. 

Sideritic shale, with many large (0.6 m diameter, or up to 0.5 m x 1 .2 m) flat, septarian, sideritic mudstone nodules; 

forms a conspicuous bed on the scar just north of Ness Ruck; occasional logs of fossil wood 0.23 

Lytoceras fimbriatum (J. Sowerby) (8; CA 2787-94). 

Shale 0.76 

Lytoceras fimbriatum (J. Sowerby) (2; CA 2785-86). 

Shale, sideritic, with scattered sideritic mudstone nodules 0.08 

Lytoceras fimbriatum (J. Sowerby) (3; CA 2782-84). 

Shale, grey, with a paler and harder middle band; contains several pieces of fossil wood 1.32 

Aegoceras (Beaniceras) luridum (Simpson) (1; CA 4564). 

Subzone of Acanthopleuroceras valdani 

Sideritic shale, with occasional sideritic mudstone nodules; crosses the scar to reach low tide mark at Ness Ruck 0.13 

Liparoceras (L.) cf. heptangulare (Young & Bird) (1 ; CA 4560). 

Shale 0.18 

Scattered sideritic mudstone nodules in shale 0.08 

Shale 0.18 

Scattered sideritic mudstone nodules in shale 0.08 

Lytoceras fimbriatum (J. Sowerby) (3; CA 2779-81). 

Shale 0.30 

Scattered flat sideritic mudstone nodules in shale 0.05 

Lytoceras fimbriatum (J. Sowerby) ( 1 ; CA 2778; PI. 1, fig. 3). 

Shale; contains fossil wood 1.17 

Widely scattered sideritic mudstone nodules in shale 0.10 

Shale 1.02 

Sideritic mudstone nodules, sometimes septarian 0.15 

Liparoceras (L.) heptangulare (Young & Bird) (1; CA 4559), Lytoceras fimbriatum (J. Sowerby) (3; CA 2775-77). 

Shale; contains fossil wood 0.74 

Sideritic mudstone nodules 0.08 

Shale 2.59 



100 M.K. HOWARTH 

571 Shale, with many scattered oval sideritic mudstone nodules up to 0.15m thick 0.81 

Liparoceras (L.) heptangulare (Young & Bird) (1; C.39137), Lytoceras fimbriatum (J. Sowerby) (1; CA 211 A). 

Subzone of Tropidoceras masseanum 

570 Shale, with occasional sideritic mudstone nodules that contain gastropods and poorly preserved Crustacea 1.73 

Lytoceras fimbriatum (J. Sowerby) (3; CA 2771-73). 
569 Continuous bed of sideritic mudstone; outcrops in a gully on the scars, and forms a conspicuous datum line 

in the cliff 0.08 

Tragophylloceras loscombi (J. Sowerby) (2; CA 2761-62). 
568 Shale, in 7 or 8 pale and dark bands 1.68 

Lytoceras sp. indet. near top of bed ( 1 ; lost): Tropidoceras futtereri Spath near base of bed ( 1 ; CA 4545; PI. 7, fig. 1 1 ); 

Tropidoceras masseanum (d'Orbigny) var. rotundum (Futterer) at boundary of beds 567 and 568 (11; CA 4546-55). 

567 Shale, hard and silty; forms a slight feature on the scar 0.15 

566 Shale 0.28 

Liparoceras (L.) sp. indet. ( 1 ; lost). 

565 Shale, hard and silty; forms a slight feature on the scar 0.08 

564 Shale, with harder silty bands at the base and the middle; the higher silty band contains occasional sideritic 

mudstone nodules 0.56 

Liparoceras (L.) sp. indet. ( 1 ; lost). 

563 Sideritic mudstone nodules; contains large logs of fossil wood 0.08 

562 Shale, with occasional sideritic mudstone nodules 0.69 

Tropidoceras sp. indet. (3; CA 4556-58), Liparoceras (L.) cheltiense (Murchison) (2; C. 39138-39). 
561 Flat sideritic mudstone nodules 0.08 

Tropidoceras sp. indet. or lUptonia cf.jamesoni (J. de C. Sowerby) (2, lost). 
560.3 Shale, with shelly layers (top 0.08 m only) 0.08 

Tropidoceras futtereri Spath (1; CA 4544; PI. 7, fig. 10). 

Zone of Uptonia jamesoni 
Subzone of Uptonia jamesoni 

560.3 Shale, with shelly layers (all below the top 0.08 m) 1.32 

Tragophylloceras sp. indet. (1; CA 2770), Polymorphites bronni (Roemer) (7; CA 4224-30; PL 7, fig. 5). Uptonia lata 
(Quenstedt) (1; CA 4538) and Uptonia cf.jamesoni (J. de C. Sowerby) ( 1 ; CA 4532). 

560.2 Flat sideritic mudstone nodules: together with bed 559. forms a distinctive pair of nodule beds on the scar 0.08 

Uptonia jamesoni (J. de C. Sowerby) (1; CA 4531), Polymorphites bronni (Roemer) (2; CA 4222-23). 

560.1 Shale 0.15 

Polymorphites bronni (Roemer) ( 1 ; C A 422 1 ). 

559 Flat sideritic mudstone nodules 0.08 

Uptonia jamesoni (J. de C. Sowerby) (1; CA 4530), Uptonia sp. indet. (1; C A 4543), Polymorphites bronni (Roemer) 

(1;CA4220). 
558 Shale, with occasional sideritic mudstone nodules at about the middle of the bed 0.66 

Uptonia lata (Quenstedt) (5; CA 4533-37), Uptonia sp. indet. (4; CA 4539-42), Polymorphites bronni (Roemer) 

( 16; CA 4204-19), Tragophylloceras sp. indet. ( 1; CA 2769). 
557 Flat sideritic mudstone nodules; conspicuous in the cliff 0.13 

Uptonia cf.jamesoni (J. de C. Sowerby) ( 1 ; CA 4529). 

556.3 Shale 0.41 

Uptonia jamesoni (J. de C. Sowerby) (1; CA 4528). 

556.2 Small spherical, grey, calcareous mudstone nodules, with a few larger flat sideritic mudstone nodules 0.08 

Uptonia jamesoni (J. de C. Sowerby) (5; CA 4523-27), Polymorphites bronni (Roemer) (3; CA 4201-03). 

556.1 Shale ■ 0.71 

Polymorphites bronni (Roemer) (5; CA 4196-4200). 
555 Grey calcareous mudstone nodules mixed with larger sideritic mudstone nodules 0.10 

Uptonia jamesoni (J. de C. Sowerby) (1; CA 4522), Polymorphites polymorphus (Quenstedt) (2; CA 4316-17; PI. 7, 

fig. 3). 
554 Shale 0.76 

Uptonia jamesoni (J. de C. Sowerby) (2; CA 4520-21), Polymorphites bronni (Roemer) ( 1 ; CA 4195), Parinodiceras 

parinodum (Quenstedt) ( 1 ; C. 39142). 
553 Flat sideritic mudstone nodules mixed with a few smaller grey calcareous mudstone nodules 0.13 

Tragophylloceras sp. indet. (1; CA 2768). 

552 Shale...! 0.33 

551 Scattered sideritic mudstone nodules, mixed with smaller grey calcareous mudstone nodules, in shale 0.08 



LOWER LIAS OF ROBIN HOOD'S BAY 101 

Uptonia jamesoni (J. de C. Sowerby) (5; CA 4515-19). 

550 Shale 0.64 

Uptonia jamesoni (J. de C. Sowerby) (5; CA 4510-14), Platypleuroceras cf. brevispina (J. de C. Sowerby) 
(3; CA 4469-71). 

Subzone of Platypleuroceras brevispina 

549 Flat sideritic mudstone nodules 0.10 

548 Shale 0.36 

Parinodiceras parinodum (Quenstedt) ( 1 ; C. 39144), Platypleuroceras brevispina (J. de C. Sowerby) 

(2; CA 4467-68), Radstockiceras sp. indet. ( 1 ; CA 3761 ). 

547 Flat sideritic mudstone nodules 0.10 

546.5 Shale, with occasional small siderite mudstone nodules in the lower part 1.02 

Platypleuroceras aureum (Simpson) (7; CA 4489-95), Platypleuroceras brevispina (J. de C. Sowerby) (2, lost), 

Polymorphites sp. indet. (1; lost). 
546.4 Scattered sideritic mudstone nodules in shale 0.08 

Platypleuroceras brevispina (J. de C. Sowerby) (9; CA 4458-66), Platypleuroceras cf. aureum (Simpson) 

(1; CA 4488), Tragophylloceras sp. indet. (2; CA 2766-67). 
546.3 Shale 0.23 

Platypleuroceras brevispina (J. de C. Sowerby) (11; CA 4448-57), Platypleuroceras aureum (Simpson) 

(5; CA 4483-87), Platypleuroceras sp. indet. (8; CA 4502-09), Polymorphites trivialis (Simpson) ( 1 ; CA 4396), 

Parinodiceras parinodum (Quenstedt) (1; C. 39143). 
546.2 Sideritic mudstone nodules 0.08 

Platypleuroceras brevispina (J. de C. Sowerby) (2; CA 4446-47), Platypleuroceras aureum (Simpson) (4; CA 4479-82; 

PI. 7, fig. 6), Polymorphites trivialis (Simpson) (1; CA 4395). 
546.1 Shale 0.33 

Platypleuroceras brevispina (J. de C. Sowerby) (10; CA 4436-45), Platypleuroceras aureum (Simpson) 

(6; CA 4473-78), Platypleuroceras sp. indet. (6; CA 4496-4501), Polymorphites trivialis (Simpson) ( 1; CA 4394). 

Flat sideritic mudstone nodules: forms a minor feature in a conspicuous gully on the scar 0.15 

Platypleuroceras brevispina (J. de C. Sowerby) (1; CA 4435). 

Shale with horizons of nodules, 4.27 m thick; forms a cambered pavement on the scar, bordered on north and south by 

conspicuous gullies; might be called the 'Polymorphites Bed'. 

Shale 0.71 

Platypleuroceras brevispina (J. de C. Sowerby) (4; CA 4431-34), Polymorphites trivialis (Simpson) (13; CA 4381-93). 

Scattered flat sideritic mudstone nodules in shale 0.10 

Polymorphites trivialis (Simpson) (3; CA 4378-80). 

Shale 0.36 

Platypleuroceras obsoleta (Simpson) (1; CA 4472), Platypleuroceras brevispina (J. de C. Sowerby) 

(38; CA 4398-4430), Polymorphites trivialis (Simpson) (3; CA 4375-77), Radstockiceras buvignieri (d'Orbigny) 

(2; CA 3749-50). 

Flat sideritic mudstone nodules 0.10 

Polymorphites trivialis (Simpson) (5; CA 4370-74), Platypleuroceras brevispina (Simpson) (1; CA 4397), 

Radstockiceras sp. indet. ( 1 ; CA 3760). 

Subzone of Polymorphites polymorphus 

Shale 0.33 

Polymorphites trivialis (Simpson) (25; CA 4347-69), Tragophylloceras cf. numismale (Quenstedt) (1; CA 2760). 

Scattered flat sideritic mudstone nodules in shale 0.10 

Polymorphites trivialis (Simpson) (20; CA 4327-46), Radstockiceras sphenonotum (Monke) (2; CA 3757-58; 
PL 5, fig. 4), Radstockiceras sp. indet. ( 1 ; CA 3759). 

Shale, with a few grey calcareous mudstone nodules 0.93 

Polymorphites trivialis (Simpson) ( 1 1 ; CA 43 19-26; PL 7, fig. 7). 

Small grey calcareous mudstone nodules 0.08 

Shale 0.32 

Scattered flat sideritic mudstone nodules in shale 0.05 

Shale 0.40 

Tragophylloceras cf. numismale (Quenstedt) ( 1 : CA 2759). 

Small grey calcareous mudstone nodules 0.08 

Shale, with two horizons of small spherical grey calcareous mudstone nodules 0.71 

Continuous bed of sideritic mudstone; forms the northern boundary of a deep gully 0.15 

Shale 0.71 

Polymorphites caprarius (Quenstedt) (10; CA 4306-15), Radstockiceras sphenonotum (Monke) (2; CA 3755-56). 



102 M.K. HOWARTH 

542.4 Scattered small dark grey calcareous mudstone nodules in shale 0.04 

Epideroceras sp. indet. (2; CA 4071-72), Polymorphites caprarius (Quenstedt) (15; CA 4292-4305), Polymorphites 
trivialis (Simpson) ( 1 ; CA 43 18), Radstockiceras sphenonotum (Monke) (3; CA 3752-54), Tragophylloceras sp. indet. 
(1;CA2765). 

542.3 Shale 0.33 

Polymorphites caprarius (Quenstedt) (6; CA 4286-91). 

542.2 Scattered small grey calcareous mudstone nodules in shale 0.04 

Polymorphites caprarius (Quenstedt) (5; CA 4281-85). 

542.1 Shale 0.58 

Polymorphites caprarius (Quenstedt) (8; CA 4273-80), Radstockiceras sphenonotum (Monke) (1; CA 3751). 

541 Flat sideritic mudstone nodules 0.08 

Polymorphites caprarius (Quenstedt) (4; CA 4269-72), Tragophylloceras numismale (Quenstedt) (1; CA 2758). 

540.3 Shale 0.53 

Polymorphites caprarius (Quenstedt) (11; CA 4258-68). 

540.2 Scattered sideritic mudstone nodules in shale 0.08 

Polymorphites caprarius (Quenstedt) (6; CA 4253-57). 

540.1 Shale 0.64 

Polymorphites caprarius (Quenstedt) (14; CA 4239-52), Hyperderoceras sp. indet. (1; CA 4053). 

539 Flat or spherical sideritic mudstone nodules 0.06 

Polymorphites caprarius (Quenstedt) (2; CA 4237-38; PI. 7, fig. 9). 
538 Shale 0.81 

Polymorphites caprarius (Quenstedt) (6; CA 4231-36). 

Subzone of Phricodoceras taylori 

537 Scattered flat or spherical sideritic mudstone nodules in shale 0.08 

536 Shale 0.74 

535 Flat or spherical sideritic mudstone nodules 0.08 

534 Shale 0.86 

533 Flat sideritic mudstone nodules 0.08 

532 Shale, with occasional sideritic mudstone nodules 1.22 

531 Large flat sideritic mudstone nodules; forms the northern boundary of a gully cut by the Dungeon Hole fault 0.15 

530.3 Shale 0.91 

530.2 Grey calcareous mudstone nodules 0.05 

Phricodoceras cornutum (Simpson) (1; CA 4060). 

530.1 Shale 0.81 

Gemmellaroceras rutilans (Simpson) (2; CA 4179-80); Pinna folium (Young & Bird) abundant. 

529 Widely scattered sideritic mudstone nodules in shale; the lowest nodule bed exposed on the northern side of the 

Dungeon Hole fault 0.05 

Apoderoceras sp. indet. (1; CA 4052). 
528 Shale 0.81 

Gemmellaroceras sp. indet. (1; CA 4194). 
527 Continuous bed of sideritic mudstone; weathers red-brown, and is yellow-brown when broken; a distinctive bed forming 

a scarp face on the northern boundary of a gully 0.18 

PYRITOUS SHALE MEMBER 

526.7 Shale, with occasional masses of pyrites, and a few calcareous mudstone nodules 1 .57 

Apoderoceras sp. indet. ( 1 ; CA 405 1 ), Tragophylloceras sp. indet. ( 1 ; CA 2764). 
526.6 Septarian calcareous mudstone nodules 0.10 

Apoderoceras cf. aculeatum (Simpson) (4: CA 4032-35). 

526.5 Shale, with occasional masses of pyrites, and a few calcareous mudstone nodules 1.02 

Apoderoceras aculeatum (Simpson) (3; CA 4029-31; PI. 6, fig. 2). 

526.4 Calcareous mudstone nodules 0.06 

Apoderoceras sp. indet. (2; CA 4049-50). 

526.3 Shale, with many masses of pyrites and 'nests' of pyrites with a radiating structure 0.25 

Apoderoceras cf. aculeatum (Simpson) (1; CA 4028). 

526.2 Calcareous mudstone nodules, some with circular septarian jointing 0.05 

Apoderoceras cf. aculeatum (Simpson) (1; CA 4027). 

526.1 Shale, with a few 'nests' of pyrites, and some calcareous mudstone nodules, especially in the lower part 1.32 

Phricodoceras cornutum (Simpson) (5; CA 4055-59). Gemmellaroceas rutilans (Simpson) (2; CA 4177-78; PI. 7. 
fig. 4), Apoderoceras aculeatum (Simpson) (3; CA 4024-26), Apoderoceras sp. indet. ( 1 ; CA 4048). 

525 Large flat sideritic mudstone nodules, with a very irregular top surface which weathers dark red; a very conspicuous bed 

crossing the scar south of the Dungeon Hole fault (see bed 520.4) 0.10 



LOWER LIAS OF ROBIN HOOD'S BAY 103 

Apoderoceras aculeatum (Simpson) ( 1 ; CA 4023), Phricodoceras comutum (Simpson) ( 1 ; CA 4070; PI. 7, fig. 2). 

524.3 Shale, with a few calcareous or sideritic mudstone nodules, and some 'nests' of pyrites 0.51 

Phricodoceras cf. comutum (Simpson) ( 1 ; CA 4054), Apoderoceras aculeatum (Simpson) ( 1 ; CA 4022; PI. 7, fig. 1 ). 

524.2 Sideritic mudstone nodules; some pyrites 0.04 

524.1 Shale 0.52 

Phricodoceras cf. taylori (J. de C. Sowerby) ( 1 ; CA 4069), Apoderoceras sp. indet. ( 1 ; CA 4047). 

523 Flat sideritic mudstone nodules 0.06 

Apoderoceras aculeatum (Simpson) (1; CA 4021). 

522.3 Shale, with occasional sideritic mudstone nodules 0.74 

522.2 Flat sideritic mudstone nodules, with circular septarian jointing 0.04 

522.1 Shale, with occasional sideritic mudstone nodules, some with viens of pyrites, and some 'nests' of pyrites 0.66 

Apoderoceras subtriangulare (Young & Bird) ( 1 ; CA 4020). 

521 Flat sideritic mudstone nodules, with circular septarian jointing 0.05 

Tragophylloceras numismale (Quenstedt) (1; CA 2757). 
520.7 Shale 0.48 

Apoderoceras subtriangulare (Young & Bird) (2; CA 4018-19; PI. 6, fig. 4). 

520.6 Sideritic mudstone nodules 0.07 

Apoderoceras subtriangulare (Young & Bird) (2; CA 4016-17). Tragophylloceras cf. numismale (Quenstedt) 
(1;CA2756). 

520.5 Shale, with many scattered sideritic mudstone nodules 0.36 

Apoderoceras subtriangulare (Young & Bird) (1; CA 4015), Apoderoceras sp. indet. (4; CA 4043-46), Phricodoceras cf. 
nodosum (Quenstedt) (1; CA 4061), Gemmellaroceras sp. indet. (2; CA 4192-93), Tragophylloceras numismale 
(Quenstedt) (4; CA 2752-55). 

520.4 Sideritic mudstone nodules, with an irregular top surface; similar to bed 525 about 50 m to the north, 

but less conspicuous 0.10 

Apoderoceras sp. indet. ( 1 ; CA 4042). 

520.3 Shale, with many sideritic mudstone nodules 0.82 

Apoderoceras subtriangulare (Young & Bird) (1: CA 4014), Tragophylloceras numismale (Quenstedt) (3; CA 2749-51). 

520.2 Sideritic mudstone nodules 0.04 

Tragophylloceras numismale (Quenstedt) (1; CA 2748). 

520.1 Shale 0.64 

Apoderoceras subtriangulare (Young & Bird) (3; CA 401 1-13), Tragophylloceras numismale (Quenstedt) (1; CA 2747). 

519 Clay; forms a conspicuous parting in the cliff, but difficult to recognize on the scars 0.03 

518 Band of sideritic mudstone, with irregular top surface; outcrops at the foot of a low scarp face that forms the northern 

side of a gully 0.04 

517.7 Shale 0.66 

Apoderoceras subtriangulare (Young & Bird) (1; CA 4010), Tragophylloceras numismale (Quenstedt) (1; CA 2746). 

517.6 Flat sideritic mudstone nodules 0.05 

517.5 Shale 0.15 

517.4 Sideritic mudstone nodules 0.05 

517.3 Shale 0.23 

Gemmellaroceras sp. indet. (2; CA 4190-91). 

517.2 Sideritic mudstone nodules 0.05 

Gemmellaroceras sp. indet. (1; CA 4189). 

517.1 Shale 0.23 

516 Large flat sideritic mudstone nodules 0.06 

515.7 Shale 0.26 

515.6 Sideritic mudstone nodules 0.08 

515.5 Shale 0.26 

Tragophylloceras cf. numismale (Quenstedt) ( 1 ; CA 2745). 

515.4 Occasional sideritic mudstone nodules in shale 0.08 

515.3 Shale 0.26 

515.2 Flat sideritic mudstone nodules 0.08 

515.1 Shale 0.26 

514 Flat sideritic mudstone nodules 0.04 

513.7 Shale 0.58 

Apoderoceras sp. indet. (1; CA 4041 ). 

513.6 Sideritic mudstone nodules 0.08 

Apoderoceras sp. indet. (1; CA 4040). 

513.5 Shale, with a few sideritic mudstone nodules 0.13 

513.4 Sideritic mudstone nodules 0.10 

513.3 Shale 0.08 

513.2 Occasional sideritic mudstone nodules in shale 0.05 



104 M.K. HOWARTH 

513.1 Shale, with occasional sideritic mudstone nodules 0.33 

Apoderoceras subtriangulare (Young & Bird) (1; CA 4009). 

512 Scattered flat sideritic mudstone nodules in shale 0.04 

511 Shale 0.25 

Apoderoceras cf. subtriangulare (Young & Bird) (1; CA 4008). 

510 Band of elongated sideritic mudstone nodules up to 1 m long, with two sets of joints at 60° 0.04 

509.3 Shale 0.18 

509.2 Scattered sideritic mudstone nodules in shale 0.08 

509.1 Shale 0.53 

Apoderoceras cf. subtriangulare (Young & Bird) ( 1 ; CA 4007), IGemmellaroceras sp. indet. ( 1 ; CA 41 88). 

508 Irregular, septarian sideritic mudstone nodules, set in pale silty shale 0.10 

507.3 Shale 0.43 

507.2 Scattered sideritic mudstone nodules in shale 0.05 

Phricodoceras cf. taylori (J. de C. Sowerby) (1; CA 4068). 

507.1 Shale, with a few sideritic mudstone nodules 0.30 

Apoderoceras subtriangulare (Young & Bird) (3; CA 4004-06). 

506 Sideritic mudstone nodules, set in pale silty shale 0.08 

Apoderoceras subtriangulare (Young & Bird) (4; CA 4000-03), Tragophylloceras sp. indet. (1; CA 2763). 

505.3 Shale .'. 0.64 

Apoderoceras subtriangulare (Young & Bird) (2; CA 3998-99). 

505.2 Shale, with many scattered sideritic mudstone nodules 0.20 

Apoderoceras subtriangulare (Young & Bird) (2; CA 3996-97). Radstockiceras buvignieri (d'Orbigny) (1; CA 3748), 
Tragophylloceras numismale (Quenstedt) ( 1 ; CA 2744; PI. 1 , fig. 1 ). 

505.1 Shale 1.30 

Apoderoceras subtriangulare (Young & Bird) (2; CA 3994-95), Gemmellaroceras tubellum (Simpson) (1; CA 4176), 
Cenoceras striatum (J. Sowerby) (1; CN 87). 

504 Sideritic mudstone nodules set in pale silty shale 0.08 

Phricodoceras taylori (J. de C. Sowerby) ( 1 ; CA 4067), Apoderoceras subtriangulare (Young & Bird) (2; CA 3992-93). 
503 Shale; 0.51 m thick in Wine Haven 0.41 

IGemmellaroceras sp. indet. (6; CA 4182-87). 
502 Occasional grey calcareous mudstone nodules, set in pale silty shale 0.08 

Phricodoceras cf. taylori (J. de C. Sowerby) (3; CA 4064-66), Apoderoceras subtriangulare (Young & Bird) 

(6; CA 3986-91; PI. 6, fig. 5). 

501.3 Shale 1.52 

Phricodoceras cf. taylori (J. de C. Sowerby) (2; CA 4062-63), Apoderoceras sp. indet. (5: CA 4036-39), 
Gemmellaroceras sp. indet. (4; CA 4181 ). 

501.2 Occasional grey calcareous mudstone nodules in shale 0.05 

Apoderoceras cf. subtriangulare (Young & Bird) (3; CA 3983-85), Gemmellaroceras tubellum (Simpson) 

(63; CA 41 13-75). 
501.1 Shale; with calcareous mudstone nodules, 0.05 m thick, at about the middle, which is the highest nodule bed on the 

foreshore on the west side of the Peak Fault complex in Wine Haven 1.83 

Bifericeras donovani Dommergues & Meister (18; CA 3793-3810; PI. 8, fig. 3), Apoderoceras subtriangulare 
(Young & Bird) (3; CA 3980-82; PI. 5, fig. 8). 

Zone of Echioceras raricostatum 
Subzone of Paltechioceras aplanatum 

500 Flat sideritic mudstone nodules; forms the north-western boundary of The Landing at Bay Town 0.08 

499 Shale, with a few large calcareous mudstone nodules; forms the north-western part of the floor of The Landing at Bay 

Town; 1.83 m thick in Wine Haven 1.57 

Paltechioceras tardecrescens (Hauer) (2; CA 4607-08), Eoderoceras armatum (J. Sowerby) (7; CA 3885-91; PI. 8, 
fig. 2), Gleviceras guibalianum (d'Orbigny) ( 1 ; CA 4606). 

498 Shale, with numerous septarian sideritic mudstone nodules; runs down the middle of The Landing 0.23 

Paltechioceras tardecrescens (Hauer) (72; CA 3573-3643; PL 4. fig. 6). Eoderoceras armatum (J. Sowerby) 
(8; CA 3878-84), Gemmellaroceras tubellum (Simpson) (15; CA 4098-41 12). 

497 Shale, dark grey with 3 paler stripes of silty shale; contains at least one log of fossil wood 2 m long; forms the south- 
eastern part of the floor of The Landing at Bay Town 2.29 

Paltechioceras tardecrescens (Hauer) (154; CA 3428-3572; PI. 4, fig. 3), Paltechioceras regustatum (Buckman) 
(2; CA 3426-27), Eoderoceras armatum (J. Sowerby) (44; CA 3834-77). Gleviceras guibalianum (d'Orbigny) 
(6; CA 3735-40), Gemmellaroceras tubellum (Simpson) (24; CA 4074-97). 

SILICEOUS SHALE MEMBER 

496 Hard calcified, silty shale; forms the capping to Landing Scar at Bay Town 1.40 



LOWER LIAS OF ROBIN HOOD'S BAY 105 

Gleviceras guibalianum (d'Orbigny) (2; CA 3733-34), Paltechioceras regustatum (Buckman) (16; CA 3410-25), 
Paltechioceras sp. indet. (6; CA 3644-49). 

In Wine Haven bed 496 caps a conspicuous scar running from 130 m east of Tan Beck waterfall into the west 
side of the Peak Fault complex, where it can be divided into: 

496c Hard calcified, silty shale 0.28 m 

496b Shale, with a few large sideritic mudstone nodules 0.21 m 

496a Hard calcified, silty shale, especially hard near base 0.91 m 

Subzone of Leptechioceras macdonnelli 

Shale 0.84 

Gemmellaroceras tubellum (Simpson) (1; CA 4073), Leptechioceras cf. macdonnelli (Portlock) (1; CA 3404). 

Harder shale, with a few sideritic mudstone nodules 0.13 

Gleviceras guibalianum (d'Orbigny) (3; CA 3730-32). 

Shale 0.15 

Eoderoceras armatum (J. de C. Sowerby) (1; CA 3833). 

Harder calcified shale 0.15 

Shale 0.30 

Hard calcified shale 0.36 

Shale 0.43 

Eoderoceras armatum (J. de C. Sowerby) ( 1 ; CA 3832). 

Harder shale 0.13 

Shale, with rare sideritic mudstone nodules in Wine Haven 0.33 

Leptechioceras cf. macdonnelli (Portlock) (1; CA 3403). 

Harder shale 0.13 

Shale 0.53 

Hard calcified, silty shale; forms the capping of East Scar at Bay Town, and forms a well-marked scar in Wine Haven 

running eastwards from Tan Beck waterfall, where a few sideritic mudstone nodules occur in the top 0.20 m 1.00 

Leptechioceras aff. macdonnelli (Portlock) (3; CA 3400-02), Eoderoceras armatum (J. Sowerby) ( 1; CA 3831), 
Radstockiceras buvignieri (d'Orbigny) (4; CA 3744-47; PI. 5, fig. 1). 

Subzone of Echioceras raricostatoides 

Shale 0.86 

Paltechioceras planum (Trueman & Williams) (1; CA 3409). 

Hard calcified, silty shale; two 0.08 m partings of softer shale seen in the Wine Haven cliff 0.91 

Paltechioceras planum (Trueman & Williams) (3; CA 3406-08). 

Shale 0.38 

Paltechioceras planum (Trueman & Williams) (1; CA 3405). 

Hard calcified, silty shale; contains a few sideritic mudstone nodules in Wine Haven 0.33 

Echioceras intermedium (Trueman & Williams) (5; CA 3383-87), Eoderoceras hastatum (Young & Bird) 
(4; CA 3892-95; PI. 6, fig. 3). 

Shale 0.22 

Hard calcified, silty shale; 0.28 m in Wine Haven; caps a strong scar south of East Scar, and a conspicuous scar in 

Wine Haven where it passes from cliff to scars at the foot of Tan Beck waterfall 0.23 

Shale 0.37 

Harder shale 0.48 

Echioceras intermedium (Trueman & Williams) (2; CA 3381-82). 

Shale, with one or two slightly harder bands 1.19 

Echioceras raricostatoides Vadasz ( 1 ; CA 3399). 

Hard calcified, silty shale; forms the highest scar that passes in front of Tan Beck waterfall in Wine Haven; crosses Mill 

Beck just west of Bay Mill 0.18 

Echioceras raricostatoides Vadasz (5; CA 3394-98). 

Shale, soft, laminated; 0.88 m in Wine Haven 0.91 

Echioceras raricostatoides Vadasz (8; CA 3389-93; PI. 4, fig. 2). 

Hard calcified, silty shale; has a softer central part and is 0.22 m thick in Wine Haven; crosses Mill Beck beside 

Bay Mill 0.15 

Echioceras raricostatoides Vadasz (1; CA 3388), Crucilobiceras densinodulum Buckman (1; CA 3830). 

Subzone of Crucilobiceras densinodulum 

Shale, with two 0.09 m slightly harder bands 0.92 

Crucilobiceras densinodulum Buckman (2; CA 3829) in lower part. 



106 M.K. HOWARTH 

486.3 Hard calcified, silty shale, with many small calcareous mudstone nodules; the 'Crucilobiceras Bed' 0.08 

Crucilobiceras densinodulum Buckman ( 18; CA 3812-28; PI. 6, fig. 1 ). 

Zone of Oxynoticeras oxynotum 
Subzone of Oxynoticeras oxynotum 

486.2 Shale 0.06 

Bifericeras cf. vitreum (Simpson) ( 1 ; CA 381 1 ). 

486.1 Hard calcified, silty shale; 0.33 m in Wine Haven, where there is a softer central 0.10m parting 0.25 

485.3 Shale; a few lenticles of calcareous mudstone in Wine Haven 0.55 

485.2 Harder shale; 0.17 m in Wine Haven 0.22 

Gleviceras doris (Reynes) ( 1; CA 3727). Angulaticeras sp. indet. (1; CA 2801 ). 

485.1 Shale 0.43 

484 Hard calcified, silty shale, with a central parting about 0.09 m thick; similar to, but less conspicuous than, bed 474, the 

Double Band; 0.56 m thick in Wine Haven 0.44 

Gleviceras cf. guibalianum (d'Orbigny) (2; CA 3728-29). 
483.5 Shale 0.13 

483.4 Harder shale 0.46 

483.3 Shale, with a few harder lenticles near the base in Wine Haven 0.43 

Oxynoticeras sp. indet. ( 1: CA 3722), IGleviceras sp. indet. (1; CA 3743). 

483.2 Harder shale; 0.09 m in Wine Haven 0.48 

483.1 Shale; 0.25 m in Wine Haven 0.18 

IGleviceras sp. indet. ( 1 ; CA 3742), Bifericeras bifer (Quenstedt) (5; CA 3788-92). 

482.5 Hard calcified, silty shale; forms the capping to the main Dab Dumps scar; 0.08 m in Wine Haven 0.10 

482.4 Shale 0.10 

IGleviceras sp. indet. (1; CA 3741). 

482.3 Hard calcified, silty shale; 0.10 m in Wine Haven 0.08 

Oxynoticeras sp. indet. (1; CA 3721). 

482.2 Shale; 0.10 m in Wine Haven 0.08 

482. 1 Hard calcified, silty shale; caps a fairly prominent scar on Dab Dumps; crosses Mill Beck nearly opposite the foot of 

Mill Bank where the road reaches the shore 0.13 

Oxynoticeras sp. indet. ( 1 ; CA 3720). 
481 Shale; 0.43 m in Wine Haven 0.38 

Oxynoticeras oxynotum (Quenstedt) ( 1 ; CA 37 16; PI. 4, fig. 4). 
480 Hard calcified, silty shale; 0.89 m in Wine Haven, where the upper half is less hard 0.80 

Oxynoticeras sp. indet. (2; CA 3718-19). 
479 Shale 0.20 

Oxynoticeras cf. oxynotum (Quenstedt) (1; CA 3715). 

478 Hard calcified, silty shale 0.14 

477 Shale, with some thin slightly harder bands 0.42 

476 Hard calcified, silty shale; 0.14 m in Wine Haven; crosses Mill Beck just inside the mouth of the valley 0.10 

Gleviceras doris (Reynes) (2; CA 3725-26; PI. 4, fig. 7). 

475.6 Shale, slightly indurated in places 0.13 

475.5 Hard calcified mudstone lenticles 0.02 

475.4 Shale, slightly harder in basal 0.05 m 0.77 

475.3 Blue or red-weathering calcareous mudstone nodules 0.05 

Oxynoticeras oxynotum (Quenstedt) (1; CA 3714). 

475.2 Shale, slightly harder in top 0.13 m 0.43 

475.1 Harder shale; 0.10 m in Wine Haven 0.08 

474 The Double Band. Hard calcified, silty shale, with a softer central parting, forming a conspicuous double band; caps 

Cowling Scar in the middle of the bay, forms a terrace in front of Mill Beck Nab and Boggle Hole, and caps Billet Scar 
in Wine Haven in the south of the bay. 

474.3 Hard calcified, silty shale; 0.25 m in Wine Haven, where there are some patches of pyrites 0.20 

IParoxynoticeras salisburgense (Hauer) (2; CA 3723-24). 

474.2 Shale, softer, only partially calcified 0.08 

474.1 Hard calcified, silty shale 0.15 

473 Shale, soft, but a partially calcified band in the middle 0.25 

Oxynoticeras sp. indet. ( 1 ; lost). 

472.3 Partly calcified shale 0.13 

472.2 Hard calcified, silty shale 0.13 

472.1 Shale, slightly calcified in places 0.61 

Oxynoticeras oxynotum (Quenstedt) ( 1 ; CA 37 1 3 ). 



LOWER LIAS OF ROBIN HOOD'S BAY 107 

Subzone of Oxynoticeras simpsoni 

471 Shale, with occasional calcareous mudstone 'cheese' doggers and masses of pyrites; 0.91 m near Miller's Nab 0.86 

470 Calcareous mudstone nodules enveloped in cone-in-cone structures, some 'cheese' doggers and a few strings of pyrites .. 0.08 

Gagaticeras exortum (Simpson) ( 1; CA 3297), Gagaticeras neglectum (Simpson) (35; CA 3348-64), Oxynoticeras 
simpsoni (Simpson) (2; CA 3711-12). 

469 Shale, with scattered calcareous mudstone 'cheese' doggers 1.83 

468 Shale, with many calcareous mudstone nodules about 0.08 m thick enveloped in cone-in-cone structures, and thin 

lenticles of shelly limestone 0.13 

Gagaticeras exortum (Simpson) (5; CA 3292-96; PI. 2, fig. 7), Gagaticeras finitimum Blake (3; CA 3306-08), 
Gagaticeras neglectum (Simpson) (28; CA 3325-47; PL 2, fig. 6), Gagaticeras sp. indet. (10; CA 3372-80), Oxynoticeras 
simpsoni (Simpson) (22; CA 3689-3710; PL 4, fig. 5), Cenoceras striatus (J. Sowerby) (1; CN 93). 

467 Calcified shale 0.10 

Oxynoticeras simpsoni (Simpson) (29; CA 3660-88), Gagaticeras exortum (Simpson) (9; CA 3283-91), Gagaticeras 
finitimum Blake (8; CA 3298-3305), Gagaticeras gagateum (Young & Bird) (2; CA 3309-10), Gagaticeras neglectum 
(Simpson) ( 14; CA 331 1-24). Gagaticeras sp. indet. (7; CA 3365-71), Palaeoechioceras sp. indet. (3; CA 3280-82). 

466 Shale, with two thin partially calcified bands; 0.48 m thick between Peter White Cliff and Miller's Nab 0.57 

Oxynoticeras cf. simpsoni (Simpson) (2; CA 3658-59). 

465 Hard sandy and micaceous calcified shale 0.08 

Oxynoticeras cf. simpsoni (Simpson) (2; CA 3656-57). 

464.33 Shale, with indurated patches and scattered 'cheese' doggers of lenticular limestone up to 0.2 m thick and up to 

2.5 m diameter 0.39 

Oxynoticeras simpsoni (Simpson) ( 1 ; CA 3655), Oxynoticeras sp. indet. ( 1 ; CA 3717), Angulaticeras sp. indet. 
(1; CA 2800), Cymbites sp. indet. (1; CA 3783). 

464.32 Band of irregularly-shaped limestone nodules in a shale matrix 0.35 

Oxynoticeras simpsoni (Simpson) (3; CA 3652-54; PL 4, fig. 8), Eparietites impendens (Young & Bird) 
( 14; CA 3262-75), Cymbites sp. indet. ( 1 ; CA 3782), Cenoceras striatus (J. Sowerby) ( 1 ; CN 86). 

464.31 Shale 0.10 

Eparietites impendens (Young & Bird) (2; CA 3260-61). 

464.2 Harder calcified shale 0.06 

464.1 Shale, with scattered calcareous mudstone nodules 1.12 

Eparietites impendens (Young & Bird) (13; CA 3248-59), Cymbites laevigatus (J. de C. Sowerby) (1; CA 3776), 
Angulaticeras sp. indet. (1; CA 2799). 

463 Calcareous mudstone nodules; many Cardinia 0.05 

Oxynoticeras simpsoni (Simpson) (2; CA 3650-51 ), Eparietites impendens (Young & Bird) (3; CA 3245-47). 

Zone of Asteroceras obtusum 
Subzone of Eparietites denotatus 

462 Shale, with occasional sideritic mudstone 'cheese' doggers; 0.91 m thick between Peter White Cliff and Miller's Nab 0.97 

Eparietites impendens (Young & Bird) (16; CA 3229-44; PL 4, fig. 1 ), Cymbites laevigatus (J. de C. Sowerby) 
(7; CA 3769-75), Angulaticeras sp. indet. (2; CA 2797-98). 'Tinkler's Stone' is a boulder of very hard grey-brown 
massive limestone, not derived from the Lower Lias, resting on bed 462 about 150 m north of the mouth of Stoupe 
Beck; it measures 1.8 m x 1.3 m x 0.85 m high and weighs about 6000 kg. 

461 Partly calcified shale 0.05 

Eparietites impendens (Young & Bird) (3; CA 3226-28; PL 1, fig. 6), Angulaticeras sp. indet. (1; CA 2796). 

460 Shale 0.29 

Eparietites cf. impendens (Young & Bird) (2; CA 3224-25). 

459 Partly calcified shale, with occasional calcareous mudstone nodules in the lower part 0.10 

Eparietites impendens (Young & Bird) (2; CA 3222-23). 

458.3 Shale 0.18 

Aegasteroceras crassum Spath (2; CA 305 1-52), Aegasteroceras sp. indet. (2; CA 3177-78), Eparietites impendens 

(Young & Bird) ( 1 ; CA 322 1 ), Cymbites sp. indet. ( 1 ; CA 378 1 ). 

458.2 Partly calcified shale 0.23 

Aegasteroceras crassum Spath (4; CA 3047-50; PL 2, fig. 4), Aegasteroceras sp. indet. (1; CA 3176). 

458.1 Shale; with occasional calcareous mudstone nodules 0.20 

Aegerlaevis (Blake) (Decapod crustacean; Withers, 1933); Aegasteroceras sagittarium (Blake) (4; CA 3172-75). 
457 Calcified shale 0.08 

Aegasteroceras sagittarium (Blake) (4; CA 3168-71), Eparietites impendens (Young & Bird) (1; CA 3220). 
456 Shale, with scattered small calcareous mudstone nodules in the lower half 0.66 

Aegasteroceras crassum Spath (1; CA 3046), Aegasteroceras sagittarium (Blake) (60; CA 3108-67; PL 1, fig. 7), 

Cymbites sp. indet. (2; CA 3779-80). 



108 M.K. HOWARTH 

455 Hard calcified shale, with two softer partings; bed 455 caps High Scar north of Stoupe Beck, and it also forms the 

highest scar in front of Miller's Nab, west of Wine Haven; the dip slope of beds 455.1 and 455.2 is pierced by 
several small excavated pools known as Strickland's Dumps (after Sir Charles Strickland). 285 m north of the mouth 
of Stoupe Beck. 

455.5 Hard calcified shale, with occasional small calcareous mudstone nodules near the top 0.15 

Aegasteroceras crassum Spath (1; CA 3045). Aegasteroceras sagittarium (Blake) (1; CA 3107), Eparietites bairstowi sp. 
nov. (2; CA 3218-19; PI. 2, fig. 8). 

455.4 Shale 0.23 

Asteroceras cf blakei Spath (2; CA 3008-09), Cymbites laevigatas (J. de C. Sowerby) (2; CA 3767-68). 

455.3 Partially calcified shale 0.10 

Asteroceras blakei Spath (4; CA 3004-07). 
455.2 Shale 0.13 

Asteroceras blakei Spath ( 1 ; CA 3003), ICymbites sp. indet. ( 1 ; CA 3778), Eparietites bairstowi sp. nov. 

(1;CA3217;P1. 3). 

Subzone of Asteroceras stellare 

455.1 Partially calcified shale 0.10 

Aegasteroceras sagittarium (Blake) (4; CA 3 103-06), Asteroceras cf. blakei Spath ( 1 ; CA 3002). 

454.2 Shale with occasional calcareous mudstone nodules 1.04 

Aegasteroceras sagittarium (Blake) (49; CA 3054-3102), Cymbites laevigatus (J. de C. Sowerby) (1; CA 3766). 

454.1 Shale with many small calcareous mudstone nodules; more calcified in lower part 0.10 

Promicroceras planicosta (J. Sowerby) (31; C.49425-31, CA 3953-75), Asteroceras blakei Spath (4; CA 2999-3001), 
Aegasteroceras sagittarium (Blake) ( 1 ; CA 3053), Cymbites laevigatus (J. de C. Sowerby) ( 1 ; CA 3765). 

453.3 Calcified shale 0.25 

Promicroceras planicosta (J. Sowerby) (10; C.49418-21, C.49423-24, CA 3950-5 \), Xipheroceras sp. indet. (1; C.49422). 

453.2 Shale 0.13 

ICymbites sp. indet. (1; CA 3777), Promicroceras planicosta (J. Sowerby) (18; C.49406-14, C.49417. CA 3944-49), 
Xipheroceras sp. indet. (1; C.49405), Asteroceras blakei Spath (3; CA 2996-98). 

453.1 Calcified shale, with many calcareous mudstone nodules 0.08 

Promicroceras planicosta (J. Sowerby) (12; C. 49402-03, C.49415-16. CA 3936-43), Xipheroceras ziphus (Zieten) 

(1; C.49404), Asteroceras sp. indet. (4; CA 3040-43), Asteroceras blakei Spath (2; CA 2994-95). 

452 Shale 0.43 

Xipheroceras ziphus (Zieten) (2; C.49400-01 ). Promicroceras planicosta (J. Sowerby) (27; C. 49383-99), Asteroceras 
blakei Spath ( 12; CA 2982-93; PI. 2, fig. 2). 

451 Calcified shale, with occasional small calcareous mudstone nodules 0.10 

Promicroceras planicosta (J. Sowerby) ( 158; C. 49369-82, CA 3918-35; PL 5, fig. 3), Xipheroceras ziphus (Zieten) 
(2; CA 3784-85; PI. 5, fig. 5), Xipheroceras sp. indet. (1; C.49368), Cymbites laevigatus (J. de C. Sowerby) 
(1; CA 3764), Asteroceras stellare (J. Sowerby) (7: CA 3028-34). Asteroceras sp. indet. (2; CA 3038-39). 

450.3 Partially calcified shale; the top 0.3 m of this bed is the lowest horizon present in the cliff; it occurs in the base of Peter 
White Cliff, and all lower beds outcrop on the wave-cut platform of the foreshore only 0.91 

450.2 Shale, with many calcareous mudstone nodules and a few 'cheese' doggers 0.10 

Asteroceras stellare (J. Sowerby) ( 1 ; CA 3027). 

450.1 Shale, with some partly calcified lenticles and a few red-weathering 'cheese' doggers 0.82 

Asteroceras stellare (J. Sowerby) (2; CA 3025-26), Promicroceras planicosta (J. Sowerby) (1; CA 3917), both species 

0.3 m above the base. 
449 Hard calcified silty shale, with well-marked jointing in two directions; forms the capping to Middle Scar between Stoupe 

Beck and Mill Beck; 0.19 m thick near Miller's Nab 0.15 

Asteroceras stellare (J. Sowerby) (7; CA 3018-24), Aegasteroceras crassum Spath (1; CA 3044). 

448.5 Shale; 0.73 m on Miller's Nab scars 0.56 

Promicroceras planicosta (J. Sowerby) (2; CA 3915-16), Asteroceras sp. indet. (1; CA 3037). 

448.4 Harder, partially calcified silty shale 0.10 

Asteroceras sp. indet. (1; CA 3036). 

448.3 Shale; 0.41 m on Miller's Nab scars 0.33 

Asteroceras stellare (J. Sowerby) (1; CA 3017). 

448.2 Grey calcareous mudstone nodules 0.09 

448.1 Shale 1.52 

Promicroceras planicosta (J. Sowerby) (3; CA 3912-14), Epophioceras landrioti (d'Orbigny) (2; CA 3277-78), 
Cymbites laevigatus (J de C. Sowerby) (1; CA 3763; PI. 5, fig. 6). 
447 Hard indurated, well-jointed, silty, calcified shale; a very conspicuous bed, forming the capping to Low Scar between 

Mill Beck and Stoupe Beck, and the capping to the most prominent scar between Stoupe Beck and Miller's Nab 0.56 

Asteroceras stellare (J. Sowerby) ( 1 ; CA 3016). 



LOWER LIAS OF ROBIN HOOD'S BAY 109 

CALCAREOUS SHALE MEMBER 

Subzone of Asteroceras obtusion 

446.5 Shale, with occasional cone-in-cone enveloped grey calcareous mudstone nodules, especially in a band from 0. 10 m 

to 0.25 m below the top 0.56 

Epophioceras landrioti (d'Orbigny ) ( 1 ; CA 3276), Xipheroceras ziphus (Zieten) ( 1 ; C.49360), Cymbites laevigatas 
(J. de C. Sowerby) (1; CA 3762), Promicroceras planicosta (J. Sowerby) (20; C.49343-59, C.49366-67). 

446.4 Harder, partially calcified shale 0.15 

Epophioceras sp. indet. ( 1 ; CA 3279), Xipheroceras cf. ziphus (Zieten) ( 1 ; C. 49337), Promicroceras planicosta 
(J. Sowerby) (1;C49338). 

446.33 Shale, with widely scattered calcareous nodules 0.25 

Promicroceras planicosta (J. Sowerby) (7; C.49362-65, CA 3909-1 1 ). Asteroceras confusum Spath (3; CA 3012-14), 
Xipheroceras dudressieri (d'Orbigny) ( 1 ; C.49336; PI. 5. fig. 2), Xipheroceras sp. indet. (2; C.49361, CA 3787). 

446.32 Large scattered calcareous nodules in a shale matrix 0.15 

Promicroceras capricornoides (Quenstedt) (4; C.49340-42, CA 3908), Asteroceras confusum Spath (2; CA 3010-1 1; 
PL 1, fig. 4), Asteroceras obtusion (J. Sowerby) (1; CA 3015), Xipheroceras sp. indet. (1; CA 3786). 

446.31 Shale, finely laminated, with occasional strings and masses of pyrites 0.60 

Promicroceras capricornoides (Quenstedt) (7; C.49339, CA 3902-07), Asteroceras sp. indet. (1; CA 3035). 

Zone of Caenisites turneri 
Subzone of Microderoceras birchi 

446.2 Cone-in-cone enveloped calcareous mudstone nodules 0.15 

446.1 Shale 0.30 

Promicroceras capricornoides (Quenstedt) (1; C. 49335). 

445 Hard calcified shale 0.13 

444 Calcareous mudstone nodules 0.06 

443.3 Shale, with a few calcareous mudstone nodules 1.37 

Promicroceras capricornoides (Quenstedt) (10; C. 49327-34). 

443.2 Hard calcified shale 0.25 

Promicroceras capricornoides (Quenstedt) (1; CA 3901). 

443.1 Grey calcareous mudstone nodules 0.05 

442.2 Hard partially calcified shale 0.48 

442.1 Grey crystalline shelly limestone, weathering yellow-brown; lenticular or nodular in some places 0.08 

441.3 Shale with occasional small calcareous mudstone nodules, especially at the middle of the bed 0.30 

441.2 Harder calcified shale and shaly limestone 0.15 

Microderoceras scoresbyi (Simpson) ( 1 ; C. 49327). 

441.1 Shale, with occasional calcareous mudstone nodules 0.41 

440 Grey shaly limestone, weathering yellow-brown 0.13 

439 Grey calcareous mudstone nodules 0.05 

438 Shale 0.15 

437 Harder partially calcified shale 0.10 

Promicroceras capricornoides (Quenstedt) (3; C. 49323-25). 
436 Shale 0.61 

Promicroceras capricornoides (Quenstedt) ( 13; C.493 15-22, CA 3896-3900), Arnioceras sp. indet. ( 1 ; CA 2938). 
435 Hard calcified shale and shaly limestone 0.15 

Caenisites cf. turneri (J. de C. Sowerby) (2; CA 321 1-12), ICaenisites sp. indet. (4; CA 3213-16), ^.Arnioceras sp. indet. 

(2; CA 2937). 
434 Grey calcareous mudstone nodules 0.06 

433.3 Shale, with impersistent calcified patches and thin shaly limestones 0.91 

Microderoceras birchi (J. Sowerby) (5; C.493 14, CA 3976-79; PI. 5, fig. 7), Caenisites turneri (J. de C. Sowerby) 

(24; CA 3187-3210; PI. 2, figs 3, 5), Caenisites brooki (J. Sowerby) (4; CA 3183-86). 

Subzone of Caenisites brooki 

433.2 Harder calified shale, with limestone lenses 0.1 1 

433.1 Shale, with occasional small calcareous mudstone nodules and cone-in-cone structures 0.15 

432 Grey shaly limestone, weathering yellow-brown, nodular in the lower half, and shelly in places 0.10 

431.3 Shale, with some thin lenses and patches of shelly limestone, especially in the upper part 0.86 

Caenisites brooki (J. Sowerby) (2; CA 3181-82), Arnioceras sp. indet. (1; lost). 

431.2 Hard, grey, shelly limestone, nodular in the lower half; some masses of pyrites 0.08 

Caenisites cf. brooki (J. Sowerby) (5; CA 3179-80, 3 lost). 



110 M.K. HOWARTH 

431.1 Shale, with a few thin lenses of limestone in the upper part 0.30 

430 Hard, pyritous, shelly, grey limestone, weathering yellow-brown 0.08 

Arnioceras sp. indet. (1; lost). 

429.8 Shale, with 0.06 m thick calcareous mudstone nodules in the top half 0.15 

429.7 Flat lenses of hard, grey, crystalline limestone 0.03 

Zone of Arnioceras semicostatum 
Subzone of Euagassiceras sauzeanum 

429.64 Grey calcareous mudstone nodules 0.05 

Arnioceras semicostatum (Young & Bird) (1; CA 2913), Coroniceras (Arietites) alcinoe (Reynes) (2; CA 2803-04; 

PL l.fig. 8). 

429.63 Shale, with thin lenses of shaly limestone, especially at the top of the lower third 0.61 

429.62 Occasional grey calcareous mudstone nodules in shale 0.05 

Arnioceras semicostatum (Young & Bird) (4; CA 2909-12). 
429.61 Shale, with small sideritic mudstone nodules, and occasional much larger 'cheese' doggers up to 2 m diameter 0.61 

Arnioceras semicostatum (Young & Bird) (2; CA 2907-08). 

429.5 Grey limestone, shaly in places, weathering yellow-brown, with small nodules on its upper surface 0.08 

429.4 Shale, containing many small grey calcareous mudstone nodules 0.08 

Arnioceras semicostatum (Young & Bird) (1; lost). 

429.3 Shale, with very occasional calcareous mudstone nodules 0.36 

429.2 Thin continuous or nodular grey limestone, shaly in places 0.03 

429.1 Shale 0.18 

Arnioceras obliquecostatum (Zieten) ( 1 ; lost), Arnioceras sp. indet. (2; CA 2935-36). 

428 Shale, with lenses or a continuous band of grey shaly limestone in the top third, and calcareous mudstone nodules in 

the lower third 0.15 

Arnioceras semicostatum (Young & Bird) (5; CA 2902-06). 

427.3 Shale 0.38 

Arnioceras semicostatum (Young & Bird) (4; CA 2898-2901). 

427.2 Irregular band of hard grey shaly limestone, weathering brown 0.15 

Arnioceras semicostatum (Young & Bird) (4; CA 2895-97), Coroniceras (Arietites) sp. indet. (1; CA 2805). 

427.1 Shale, with some lenses of shaly limestone, and a few large 'cheese' doggers of sideritic mudstone 0.71 

Arnioceras sp. indet. ( 1 ; CA 2934). 

426.2 Hard grey shaly limestone, weathering brown, with a rough top surface; forms a conspicuous scar landward of Pseudo 

Low Balk (bed 424.2); many Gryphaea 0.05 

Coroniceras [Arietites) alcinoe (Reynes) (1; CA 2802), Arnioceras semicostatum (Young & Bird) (10; CA 2885-94), 
Arnioceras sp. indet. (1; CA 2933). 

426.1 Hard calcified shale, weathering brown 0.15 

Arnioceras semicostatum (Young & Bird) (3; CA 2882-84), Euagassiceras sp. indet. (1; CA 2973). 

425.7 Shale, with a few indurated patches 0.15 

425.6 Grey calcified shale, weathering brown 0.11 

Arnioceras semicostatum (Young & Bird) (9; CA 2873-81). 

425.5 Shale; very occasional large calcareous mudstone 'cheese' doggers 0.15 

Arnioceras semicostatum (Young & Bird) (16; CA 2857-72), Arnioceras miserabile (Quenstedt) (2; CA 2919-20). 

425.4 Grey calcified shale, weathering brown 0.1 1 

Arnioceras sp. indet. ( 1 ; CA 2932). 

425.3 Shale 0.15 

Arnioceras sp. indet. (1; CA 2931). 

425.2 Grey calcified shale, weathering brown 0.1 1 

Arnioceras sp. indet. ( 1 ; CA 2930). 

425.1 Shale, with a few indurated patches 0.52 

Euagassiceras sp. indet. (2; CA 2971-72), Arnioceras sp. indet. (11; CA 2921-29). 

424.3 Calcified shale, harder in the upper half; occasional small calcareous mudstone nodules 0.61 

Euagassiceras sp. indet. (3; CA 2968-70), Arnioceras semicostatum (Young & Bird) (18; CA 2839-56), Arnioceras 
miserabile (Quenstedt) (3; CA 2916-18), Coroniceras (Arietites) alcinoe (Reynes) (1; C.41310). 

424.2 Shale, partially calcified in the top half; this middle division of bed 424 forms the scar 'Pseudo Low Balk' which 

emerges immediately landward of Low Balk (bed 422.2) 0.30 

Arnioceras semicostatum (Young & Bird) (10; CA 2829-38; PI. 1, fig. 2), Arnioceras miserabile (Quenstedt) 
(2; CA 2914-15). 

424.1 Shale, partially calcified near the top 0.61 

423 Shale, no subdivisions observed 1.58 

422.2 Hard blue-grey flaggy limestone, weathering brown, with a rough top surface; slightly micaceous; forms the capping of 



LOWER LIAS OF ROBIN HOOD'S BAY 111 

the very conspicuous scar Low Balk 0.10 

Euagassiceras resupinatum (Simpson) (21; CA 2942-62; PI. 1, fig. 5). 

422.1 Hard grey micaceous calcified shale, weathering brown 0.30 

Euagassiceras resupinatum (Simpson) ( 1; CA 2941 ). 

421.4 Hard partially calcified shale, micaceous and with strings of pyrites in places; a few harder calcified lenses near the top .. 0.91 

Arnioceras semicostatum (Young & Bird) (31; CA 2806-28, CA 2974-81), Euagassiceras resupinatum (Simpson) 
(2; CA 2939-40), Euagassiceras sp. indet. (5; CA 2963-67). 

421.3 Shale; forms a terrace on the scar due to a harder band at the top 0.46 

421.2 Shale; forms a terrace due to a harder band at the top 0.69 

Euagassiceras sp. indet. ( 1 ; lost). 

421.1 Shale; forms a terrace due to a harder band at the top 0.91 

Euagassiceras sp. indet. (2; lost), Arnioceras sp. indet. (1; lost). 
420 Hard grey flaggy limestone passing down into grey shale; slightly micaceous and pyritous; full of bivalves in places; 

forms a dip slope 25 m wide and a scarp face 0. 13 m high 1 .75 

419 Hard grey calcified shale, slightly micaceous; dip slope 1 m wide, and scarp face 0.13 m high 0.13 

418 Hard grey partially calcified shale; slightly micaceous; forms a terrace 4 m wide, ending seaward in a scaip face 0.46 m 

high; the lowest bed exposed at low water of the lowest spring tides 0.60 



LITHOSTRATIGRAPHY 



The Lower Lias of Robin Hood's Bay is 1 63.74 m thick and belongs 
to the Redcar Mudstone and the lower part of the Staithes Sandstone 
Formations. These names were introduced as formations by Powell 
(1984: 53) and Howard (1985: 262), and details of their definitions 
were given by Cox et al. (1998: 35,39). Fig. 18 is a complete vertical 
section of the succession of the Lower Lias in Robin Hood's Bay, 
showing the changes in lithology, the lithological divisions and the 
main features of significance formed by the harder beds, some of 
which have received formal names; bed thicknesses are drawn to 
scale on this figure, giving a visual indication of the relative thick- 
nesses of the subzones and lithostratigraphical divisions. 



Staithes Sandstone Formation (beds 591-601 
higher) 



and 



Consists of sandstones, that are mid to pale grey, fine to medium- 
grained, micaceous, with grey siltstone bands and some beds of silty 
shales; nodules of argillaceous limestone occur in the shales and are 
occasionally sideritic; there is much bioturbation, cross-bedding and 
ripple marked bedding, especially in the sandstones. 

The type area is in Robin Hood's Bay. The formation is somewhat 
transitional from the underlying Redcar Mudstone Formation, but a 
convenient marker bed that defines the lower boundary just south of 
Castle Chamber is bed 591, the Oyster Bed, which is a hard, 
calcified, silty shale or argillaceous sandstone, containing many 
oysters and other fossils. Above this level the amounts of silt and 
sand are higher than lower in the sequence and hard and soft 
sandstones are frequent. The formation has no subdivisions and 
extends up into the lower half of the Upper Pliensbachian. The 
Lower Pliensbachian (ie. Capricornus and Figulinum Subzones) part 
of the formation is 12.74 m thick. 



Redcar Mudstone Formation (beds 418-590; 151 
thick) 



m 



Consists of mudstones and shales, grey, soft and well-bedded, but 
some beds are indurated due to calcification, and there are some 
harder siltstones; thin beds of shelly limestone occur in the lower 
part, and nodules of calcareous or sideritic mudstone occur, especially 



in the upper part; pyritized nodules or irregular aggregations of iron 
pyrites occur at some horizons. 

Although the name is derived from the Lower Lias exposures at 
Redcar to the north-west, and the lower boundary is defined in the 
BGS Felixkirk Borehole (Cox et al. 1998: 35), the type section 
consists of the whole sequence exposed in Robin Hood's Bay below 
the base of the Staithes Sandstone Formation. Here the lithology is 
more argillaceous than in the Staithes Sandstone Formation, though 
different parts are variously more calcareous, siliceous, pyritous or 
ferruginous, and they form the basis of the following four members: 

4. Ironstone Shale Member. 
3. Pyritous Shale Member. 
2. Siliceous Shale Member. 
1 . Calcareous Shale Member. 

These members were introduced as 'Shales' by Buckman (1915: 
61 ) for lithological divisions in Robin Hood's Bay below the 'Sandy 
Series' (= Staithes Sandstone Formation), and he based them on 
groups of the ammonite zones that he applied to the succession. His 
zones can be linked to beds in his detailed sections, but they are not 
the same as the ammonite zones recognized now, and in any case it 
is not satisfactory to base lithological divisions on the ranges of 
palaeontological zones. No formal definitions have been given to 
these divisions, though Hesselbo & Jenkyns ( 1 995 : 114-135) applied 
them informally to the succession, and placed boundaries between 
them at appropriate levels in the succession, all of which are followed 
here. The four members are defined formally here, and their type 
sections are in Robin Hood's Bay. 

Ironstone Shale Member (beds 527-590; 62.73 m thick). Mud- 
stones and shales, grey, soft, with some beds of micaceous silty shales, 
many grey sideritic mudstone nodules, weathering red on the outside, 
and a few calcareous mudstone nodules. The nodules are scattered 
sparsely through the shale, but are more often developed at single 
horizons, either scattered or as near-continuous beds. The distinctive 
beds of red- weathering nodules have frequently been referred to as 
'ironstones', but they are better described as sideritic mudstones. 
The base is defined at the bottom of bed 527 in Robin Hood's Bay, 
which is a prominent continuous bed of sideritic mudstone weather- 
ing red-brown. Although sideritic mudstone nodules occur at several 



112 



M.K. HOWARTH 





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Fig. 18 Vertical section of the Lower Lias of Robin Hood's Bay, showing the main lithological features, relative thickness of all the beds, and named beds 
and other beds that form prominent features on the foreshore. 



LOWER LIAS OF ROBIN HOOD'S BAY 



113 



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Low Balk 



114 



M.K. HOWARTH 



lower horizons down as far as bed 500 and those of bed 525 are 
especially conspicuous, pyrites is very common at most horizons up 
to bed 526.7 but does not occur higher. So the strong sideritic 
mudstone of bed 527 is a good base for this member. Such sideritic 
mudstone nodules and occasional continuous beds are a feature of 
the whole thickness of the member up to the base of the Staithes 
Sandstone Formation. Several of them form prominent features on 
the scars, eg. beds 527, 531, 543, 545, 559 and 560.2 (a distinctive 
pair of nodule beds), 569, 578.4 and 589. 

Pyritous Shale Member (beds 497-526; 26. 1 8 m thick). Consists 
of dark grey, soft and micaceous shales, with many calcareous and/ 
or sideritic mudstone nodules; there are many nodules or irregular 
masses of iron pyrites, especially in the lower part. This member is 
similar to the Ironstone Shale Member in containing both calcareous 
and sideritic mudstone nodules, but it also contains much iron pyrites 
as irregular nodules or masses of crystals. 

The base is defined here as the bottom of bed 497 in Robin Hood*s 
Bay. This is the horizon at which the sand and silt content almost 
disappears and calcification is much diminished, leaving the beds 
above as softer shales. Iron pyrites is common at many horizons, 
appearing variously as irregular masses or strings of pyrites, or pyrite- 
rich concretions, and many of the fossils are partly pyritized in these 
beds. The softness of the shale leads to this part of the succession 
forming the wettest and lowest part of the bay relative to sea level. 

Siliceous Shale Member (beds 447-496; 38.74 m thick). Dark- 
grey shales, interbedded with much harder and lighter-coloured beds 
of calcified mudstones, silts and fine sandstones; a few nodules and 
doggers of calcareous or sideritic mudstone occur. This member 
forms the series of hard calcified beds alternating with soft shales 
that is typical of the lower part of the succession in Robin Hood's 
Bay. Red-weathering sideritic nodules are now scarce, and most of 
the harder beds are calcareous cemented muds and silts, in which the 
arenaceous content is higher than in the underlying Calcareous Shale 
Member. Large, circular 'cheese' doggers of hard argillaceous lime- 
stone, containing vertically orientated cystals of calcite, occur at 
several horizons in both this member and the Calcareous Shale 
Member below. Such doggers are generally up to only about 10 cm 
thick, but they can reach 2.5 m in diameter; they occur at 7 levels 
between beds 450 and 47 1 . 

The base is defined here at the bottom of bed 447 (Low Scar) in 
Robin Hood's Bay. This is the first prominent bed of very hard 
calcified shale or argillaceous limestone that has a significant sand 
content. Sand and silt occur in many of the hard calcified shales or 
sandstones at horizons up to the base of the Pyritous Shale Member. 
Most of the prominent 'scars' in the bay are formed of beds in this 
member - ie. Low Scar, Middle Scar (Gryphaea Scar), High Scar 
(Lower Triplet), Double Band (Cowling Scar and Billet Scar). Upper 
Triplet, East Scar and Landing Scar. 

Calcareous Shale Member (beds 41 8-446; 23.35 m thick). Dark- 
grey shales, interbedded with hard, calcified, silty mudstones; doggers 
of calcareous mudstone and some beds of limestone also occur; 
cone-in-cone enveloped calcareous mudstone nodules also occur at 
several horizons. This member is less arenaceous than the Siliceous 
Shale Member, the hard beds now having no sand and less silt, and 
the hardness being due mainly to calcification. The very prominent 
Low Balk and the less prominent Pseudo Low Balk are formed by 
limestones or highly calcified shales. Large 'cheese' doggers, simi- 
lar to those in the Siliceous Shale Member, are found at three levels 
in beds 425^129. 

In Robin Hood's Bay the base has to be placed at the lowest 



horizon exposed, ie. at the base of bed 418. If it is thought that the 
base should coincide with the base of the Redcar Mudstone Forma- 
tion, then it must be defined at the same level (ie. 288.87 m depth) in 
the BGS Felixkirk Borehole (Cox etal. 1998: 35). 



EXPOSURES IN ROBIN HOODS BAY NOW 

The extent of the foreshore exposures of the solid geology on the 
north Yorkshire coast has always depended on the vagaries of shift- 
ing sand and boulder cover, algal growth, barnacle growth, and major 
cliff falls, all caused or cleared away by the actions of tides and 
storms. But in the last 25 years much more extensive, and possibly 
more permanent, sand, boulder, algal and barnacle cover, and mussel 
beds have made major inroads into the amount of rock exposed in 
some areas. Especially serious on some of the scars are mussel beds 
that trap mud and silt to form a thick, impenetrable cover that 
completely obscures the rock underneath. At the end of the 1 990s the 
foreshore exposures were largely obscured from Way Foot at the 
bottom of Robin Hood's Bay Town northwards to just south of 
Dungeon Hole. In fact there are few or no exposures of beds 497-525 
owing to the sand and seaweed cover, which has possibly been 
exacerbated by the high concrete seawall built to protect Robin 
Hood's Bay in 1975. That seawall covers the cliff face of the same 
beds, so that they are not now exposed in either the cliff face or on the 
foreshore. Exposures improve upwards from bed 526, especially 
north of the Dungeon Hole fault, though there is still much algal 
growth and large areas are covered by loose boulders. Around the 
north side of the bay in the top half of Map 1 (Fig. 5) the foreshore 
continues to be washed clean by tides and storms and exposures are 
still good. Exposures seaward and south of Boggle Hole (Map 3; Fig. 
8) are also better, and they are good in front of Peter White Cliff (Map 
4; Fig. 1 1 ). The lowest beds on the latter map, especially from bed 
430 down to below Low Balk have always suffered from algal cover, 
of which Laminaria is a significant factor at those low sea-levels, but 
barnacle growth is also very pronounced and makes observations 
difficult on some beds. 

For most of the past two centuries the foreshore of Robin Hood's 
Bay has been largely clear of such cover, and collectors from Young 
& Bird in the 1 820s, Phillips, Simpson, Tate & Blake, the Geological 
Survey in 1880-1910, up to Bairstow in the period from 1928 to the 
1950s (see Fig. 7) were able to make significant fossil collections 
from all the beds. In particular, most of them obtained large speci- 
mens of Apoderoceras from the Taylori Subzone of beds 501 to 526. 
No such specimens can be collected today. The foreshore was largely 
clean in 1969 when Bairstow conducted a field party from the 
William Smith Jurassic Symposium to the Bay, but deterioration 
proceeded rapidly from the early 1970s. Bairstow's work could not 
be repeated today, at least for the beds on Map 2 (Fig. 6) from Robin 
Hood's Bay town northwards to the top of that map. 



CORRELATION WITH PREVIOUS 
DESCRIPTIONS 

The Lower Lias of Robin Hood's Bay was mentioned by Young & 
Bird (1822, 1828) and Phillips (1829, 1875), and a few ammonites 
from the bay were figured by them, but their descriptions were not in 
sufficient detail to be correlated with the work in this paper. Prior to 
Bairstow's work, detailed descriptions were published by Simpson 
(1868, 1884), Tate & Blake (1876) and Buckman (1915). all of 
whom numbered their beds from the top downwards. After Bairstow 
prepared his maps and stratigraphical descriptions, detailed accounts 



LOWER LIAS OF ROBIN HOOD'S BAY 



115 



of the geology of parts of the bay were published by Howarth ( 1 955), 
Phelps ( 1985) and Hesselbo & Jenkyns (1995). The tables of Figs 19 
and 20 give bed-by-bed correlation columns for all these schemes in 
as much detail as is possible; the columns of subzones in both figures 
are the subzones as determined in this paper. 

Simpson first described the beds in 1 868 (Simpson, 1 868: 53-56), 
but in his later work (Simpson, 1884: xvii-xxii) there are more 
details at some horizons, and the Simpson columns in the tables are 
based on his later work. Tate & Blake (1876: 63-65, 73-75, 79-81, 
91, 92, 109, 1 10) described the beds in greater detail, and in many 
parts of the succession their beds correspond closely with those in 
the present paper, though they omitted about 3 m of strata within 
their group Jam.30-Jam.32. Tate & Blake (1876: 79, 91-2) also 
duplicated part of the succession in their descriptions, inasmuch as 
their Jamesoni beds 1-7 are the same as their Capricornus beds 28- 
33 (these are shown in Fig. 19 as Jam.l-Jam.7 only). 

The Geological Survey's description of the succession first 
appeared in the memoir of Fox-Strang ways & Barrow (1882: 4-10), 
where the beds were described in detail but not numbered, and the 
ammonites that they listed for individual beds are not accurately 
determinable in modern terms. The same description was used by 
Buckman (1915: 67-74) in his appendix to the 2 nd edition of that 
memoir: the same basic data was used for the succession, but the 
beds were sometimes combined into thicker units and were now 
given numbers; lithological names were given to a few of the beds, 
and Buckman's determinations of the ammonites and zonal divi- 
sions were added. Buckman's 1915 description of the sequence is 
used for Figs 19 and 20, rather than the original 1882 description. 

Howarth (1955: 155) described beds upwards from the bottom of 
the Upper Pliensbachian; the equivalence of his beds 1-6 are shown 
at the top of Fig. 19, with bed 1 being the same as beds 600.5 and 
600.6, while the equivalence of a few lower beds that were given 
roman numbering in the upper part of the Figulinum Subzone is 
indicated in the detailed stratigraphical section above (p. 98). 

Phelps (1985: fig. 4) described the succession in the Davoei and 
upper part of the Ibex Zones in detail. When the vertical tabular 
section of his fig. 4 is compared at the same scale with the tabular 
section of Fig. 18 here, good correlations can be made from his top 
bed near the top of the Figulinum Subzone down to his bed 4b (= bed 
567) in the Masseanum Subzone, and at the bottom it seems fairly 
certain that his bed 1 is the same as bed 561. Phelps (1985: pi. 1, figs 
1, 3, pi. 2, figs 1, 6, 8) figured five ammonites from his beds 21, 23, 
37, 47 and 63, the identifications of which are discussed below 
(pp. 141-144) in the description of the ammonite genus Aegoceras. 

Hesselbo & Jenkyns's (1995) sequence of the Lower Lias of 
Robin Hood's Bay was based on new observations made by them. 
Their bed numbers are original from the bottom of the sequence up 
to their bed 1 2 1 at the base of the Masseanum Subzone, then higher 
up they used the bed numbers of Phelps (1985), and finally the bed 
numbers of Howarth (1955) upwards from the top of the Figulinum 
Subzone. Although their descriptions and measurements were new, 
above bed 1 2 1 their identification and use of Phelps ' bed numbers is 
difficult to interpret at some horizons, especially in the Ibex Zone, so 
in Fig. 19 the correlation of Phelps' beds 1-65 is based on Phelps' 
original description of that sequence, not on Hesselbo & Jenkyns' re- 
interpretation of it. However, from the base of the sequence up to bed 
121, Hesselbo & Jenkyn's description can be readily correlated with 
that of this paper at most levels, and is shown as their beds 1-121 in 
the relevant columns of Figs 19 and 20. The main areas of uncer- 
tainty are at the bottom of the succession below their bed 23 (=bed 
447), though it appears likely that their bed 5 has been correctly 
identified as bed 422 (Low Balk), in their beds 73-94 in the Taylori 
Subzone, that are difficult to correlate in detail, and in beds 113-121 



in the Brevispina and Jamesoni Subzones. In the upper part of the 
sequence between beds 102 and 1 16, a strikingly similar pattern can 
be seen by comparing Hesselbo & Jenkyns' tabular section side-by- 
side with that ofFig. 17;eg. bed 102 = bed527; 104 = 531; 1 12 = 543; 
1 14, lower part = 545, and it is probable that bed 1 16 is the same as 
bed 547. Bed 1 of Phelps has already been correlated with bed 561, 
so this leaves Hesselbo & Jenkyns' beds 117-121 (4.5 m thick) as 
eqivalent to Bairstow beds 548-560 (6.3 m thick), but there are some 
differences in thickness and they are not correlatable in detail. The 
position of the subzone boundaries given by both Phelps and Hesselbo 
& Jenkyns differ in detail from those determined for this paper, 
except for the upper parts of the Raricostatum and Davoei Zones. 

In addition to the previous descriptions in the works listed above, 
Getty measured and collected ammonites from the Oxynotum and 
Raricostatum Zones in the bay. The stratigraphical part of his work 
is only available in his unpublished thesis (Getty, 1972), but many of 
the ammonites he collected were described in his revision of the 
family Echioceratidae (Getty, 1973), and they are in the collections 
of the Natural History Museum. His stratigraphical sequence of 
ammonites and biostratigraphical divisions are very similar to those 
of Bairstow as determined here. 



BAIRSTOW'S AMMONITE COLLECTION 

More than 2360 ammonites were collected by Bairstow. The major- 
ity were obtained in the years 1927-1935, but small numbers of 
specimens were added up to about 1970. In addition there are a few 
specimens that were given to him by other collectors: the majority 
came from Dr J. Coggin Brown, who collected well-preserved 
ammonites at Robin Hood's Bay in the period 1940-1960 (on 
retiring to north-east England after working for the Geological 
Survey of India). Bairstow checked the horizons of the specimens 
given to him with great care, and only those that he was satisfied 
came from definitely identifiable beds are included amongst those 
listed in this paper. All Bairstow's ammonites are preserved in 
collections of the Department of Palaeontology, The Natural History 
Museum, London, and most have been given Museum registration 
numbers, in addition to the collecting numbers given by Bairstow. 
The identifiable Liparoceratidae were registered for Spath's (1938) 
catalogue of that family, and received some of the numbers in the 
series C.38871-C. 39579; some of the Eoderoceratidae were regis- 
tered in the late 1950s in the series C.493 14-C.4943 1 ; the remainder 
of the collection was registered in 2000 with the numbers CA 2744- 
CA 4608. The three nautiloids in his collection have the numbers CN 
86, 87 and 93. 

In 1928 Bairstow consulted with S.S. Buckman, a year or two 
before his death, who had expressed interest in the ammonites he was 
collecting in the bay. From his earlier work on ammonites collected 
by the Geological Survey, Buckman knew that the succession up to 
the top of the Sinemurian was exposed in both the north-western and 
south-eastern parts of the bay. The sequences of ammonites that 
Bairstow was obtaining in the two outcrops that are up to 3 km apart 
seemed to Buckman to be a good opportunity to test his hemeral 
theory 1 , and he advised Bairstow to record the geographical position, 



'Briefly, Buckman's theory of hemera was that every species of ammonite reached its 
acme of abundance at a unique time that did not overlap with the acme of any other 
species. By discovering the order in which ammonites reached their acme, a sequence 
of 'hemerae' could be constructed, which would be smaller and finer divisions than 
ammonite subzones and would be applicable over wide areas. Contemporary palaeon- 
tologists were sceptical of the theory, and work by many palaeontologists during the 
following 70 years has shown that the hemeral theory is not valid. 



116 



M.K. HOWARTH 



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588 


LLe, 


Cap.2 1 


587 


LLe 2 


Cap.22 


24 


586 


LLf, 


Cap.23 


23 


585 


LLf 2 


Cap.24 


22 


584 


LLg, 


21 


583.2 


Cap.25 


583.1 


LLg, 


Cap.26 


20 


582 


LLh, 


Cap.27 


19 


581 


LLh 2 


Jam.l 


18 


580 


E 

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3 


LLi, 


Jam. 2 




17 


579 




Jam. 3 




16 


578.5 




Jam.4 




15 


578.4 




Jam.5 






578.3 




Jam.6 




578.2 




Jam.7 




578.1 




Jam.8 






577 


'5 
a 
-a 

> 


LL i 2 -r 


Jam. 9 




13 


S76.4-.9 




12 


576.1-.3 




Jam. 10 




11 


575 




Jam. 11 




10 


574 




Jam. 12 




9 


573 




Jam. 13 




8 


572 




Jam. 14- 16 




7 


571 




Jam. 17 




6 


570 


a 




Jam. 18 


Ch.8 


5 


569 




Jam. 19 




4c 


568 


2 





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567 






4a 


564-66 


3 




Jam.20 




3 


563 


S 




Jam. 21 




2 


562 


a 




Jam.22 




1 


561 


' S 




Jam.23 




117-121 


560.3 










560.1 -.2 






Jam. 24 




559 






Jam.25 




558 


c 




Jam.26 




557 






Jam.27 


Ch.8 


556 


" E 




Jam.28 




555 


-> 




Jam. 29 


554 






Jam.30-32 


550-553 






548, 549 






115, 116 


547 


ta 

c 

Q. 
> 

CQ 




546.5 




114 


546.1 -.4 




545 




Jam.33 


113 


544.6-. 9 




544. 1 -.5 


3 
& 

o 
E 
>> 

o 
a 




LLs, 


Jam. 34 


Ch.9 


112 


543 




LLs, 


Jam.35 


111 


542 




Jam.36 


110 


541 




Jam.37 


109 


540 




Jam.38 


108 


539 




Jam. 39 


107 


538 




536, 537 






LLt, 


Jam.40 


535 




LLt 2 


Jam.4 1 


534 




LLu, 


Jam.42 


106 


533 




LL u, 


Jam.43 


105 


532 




LLv, 


Jam.44 


104 


531 




LLv 2 


Jam.45 


Ch.10 


103 


530.2-.3 




530.1 




LLw, 




LL w. 




528, 529 




LL x. 


Jam.46 


102 


527 






Jam. 47 


Ch.ll 


101 


526.7 




100 


526.6 




99 


526.5 




98 


526.2-.4 




97 


526.1 




Jam.48 


Ch.l 2 


96 


525 




Jam.49 


95 


522-24 




94 


521 




93 


520 




Jam.50 


92 


519 




Jam.5 1 


518 




Jam. 52 


91 


517 




Jam.53 




90 


516 




LL x 2 -y 2 


Jam.54, 




89 


513.7-515 






88 


513.2-.6 






87 


511-513.1 






86 


509.2-510 






85 


509.1 






84 


508 




Ch.13 


83 


507 






82 


506 






81 


505.3 






80 


505.2 






77-79 


505.1 






76 


504 








501.3-503 




Jam.54, 


Ch.14 


73-75 


501.2 


L 




Jam.55 


Ch.l 5 




501.1 



Fig. 19 Correlation of the bed numbers used in this paper for the Lower Pliensbachian with divisions used in previous descriptions of the Lower Lias of 
Robin Hood's Bay; the subzones in the right hand columns are those determined in this paper. See text for details of the sources of the previous 
descriptions. 



LOWER LIAS OF ROBIN HOOD'S BAY 



117 



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Ch. 17 


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499 


Jam. 58 


Ch. 18 


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498 


Jam. 59 


Ch. 19 


Jam. 60 


Ch. 20 


69 


497 


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Ch. 22 


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496 


LL 1,-13 


Ox. 2 


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495.7 


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495.3-.6 


65 


495.15-.2 


64 


495.11-. 14 


Ox. 3 


63 


494 


Ox. 4 


62 


493.5 


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493.4 


60 


493.1 -.3 


Ox. 5 


59 


492 


Ox. 6-12 

. . . _ 7 

Ox. 13 


58 


491 


57 


490 


56 


489 


55 


488 


54 


487 


Den. 


53 


486.3 


486.1-.2 


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485 


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484 


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483.4-.S 


49 


483.1-.3 


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482 


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480-81 


Ox. 14 


479 


Ox. 15 


46 


478 


Ox. 16 


45 


477 


Ox. 17 


44 


476 


Ox. 18 


Ch. 23 


475.4-.6 


Ox. 19 


475.3 


Ox. 20 


Sin. 1 


475.1 -.2 


LL 14, 


Ox. 21, 


Sin. 2 


43 


474.3 


Ox. 21 2 


474.2 


Ox. 22 


474.1 


LL 14 2 


Ox. 23 


Sin. 3 


42 


472.3-473 


LL15, 


Ox. 24 


472.2 


LL 15, 


Ox. 25 


472.1 


471 


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Ox. 26 


41 


470 


Ox. 27 


40 


469 


39 


468 


LL 16, 


Ox. 28 


Sin. 4 


38 


467 


LL 16, 


Ox. 29 


466.3-.5 


LL 17, 


466.2 


LL 17, 


466.1 


LL 18, 


Ox. 30 


37 


465 


LL18, 


Ox. 31 


464.3 


LL 19, 


464.2 


LL19, 


464.1 


LL20, 


36 


463 



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34 


461 


LL21, 


458-60 


Ox. 32 


457 


Ox. 33 


33 


456 


Ox. 34 


Sin. 9 


32 


455.5 


Ox. 35 


455.2-.4 


455.1 


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31 


454 


Ox. 36 


Sin. 10 


30 


453.3 


Ox. 37 


453.1-.2 


29 


452 


Ox.38, 39 


28 


450.3-451 


27 


450.1-.2 


LL22, 


Ox.40 


26 


449 


LL22, 


Ox.41 


25 


448.5 


24 


448.1 -.4 


LL23, 


Ox. 42 


Sin. 11 


23 


447 


LL 23 2 -26 


Ox. 43 


22 


446.5 


3 

O 


Ox. 44 


446.4 


Ox. 45 


446.3 


21 


446.2 


IS 

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20 


446.1 


Buckl. 1 


445 


Buckl. 2 


444 


19 


443.3 


18 


443.1-2 


17 


442.2 


Buckl. 3 


442.1 


Buckl. 4 


441 


Buckl. 5 


16 


440 


Buckl. 6 


438-39 


Buckl. 7 


437 


Buckl. 8 


436 


15 


435 


14 


433.3-434 


433.1 -.2 


o 

o 

pa 


Buckl. 9 


432 


Buckl. 10- 
21 


Sin. 12 


431.3 


13 


431.2 


12 


429.7-431.1 


429.6 


E 

3 
C 

ca 

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N 
3 
(S 
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11 


429.5 


9, 10 


429.1 -.4 


426-28 




Sin. 13 


425 




8 


424.3 


6,7 


424.1 -.2 


423 


LL27, 


5 


422.2 


422.1 


LL27,.... 


3,4 


421 


1,2 


420 




418-19 



Fig. 20 Correlation of the bed numbers used in this paper for the Sinemurian with divisions used in previous descriptions of the Lower Lias of Robin 
Hood's Bay; the subzones in the right hand columns are those determined in this paper. See text for details of the sources of the previous descriptions. 



as well as the stratigraphical horizon, of all his specimens. Bairstow 
did this by drawing 1 7 datum lines on his copies of the maps (ie. they 
are not on the originals in King's College, Cambridge). Each was a 
straight line crossing the foreshore from the base of the cliff to the 
seaward edge of the scar; most cross the foreshore approximately at 



right angles to the cliff, but a few cross at an oblique angle. Originally 
he chiselled marks on the outcrops to record the exact position of 
each datum line, but all such marks have long since disappeared. 
Each ammonite he collected was related to the nearest point on a 
datum line by pacing yards (0.9 m) from the intersection of the line 



118 



M.K. HOWARTH 



with the bottom of the cliff, then yards at right angles to the line up 
to the position of the ammonite. By these means the geographical 
position of each ammonite was recorded to approximately the near- 
est square yard. This information, which is not included in this paper, 
occurs on many of the original specimen labels that are with the 
ammonites in his collection. 

In 1957 Bairstow prepared a bed-by-bed list of the identifications 
of every ammonite in his collection. This is a large manuscript 
amounting to 390 pages. Not only is it a list of the specimens then in 
the collection (over the years a small number had decayed or were 
lost), but its main value is as a record of the identifications made by 
Dr L.F. Spath. He saw the specimens as Bairstow collected them, and 
made identifications that date mainly from the period 1927^-0, 
while a few were checked or reidentified by him up to 1956. In 
preparing the list of ammonites for this paper, all the identifications 
were verified, mainly in order to produce a consistent set of 
determinations from which the account of the biostratigraphy could 
be prepared, but also to revise the generic attributions of the species 
according to modern usage of the various genera. In general Spath's 
identifications were found to be accurate, and only a few needed 
revision. The only previous publication of any of Spath's identifica- 
tions was in his catalogue of the Liparoceratidae (Spath, 1938), 
where all the Robin Hood's Bay Liparoceratidae collected up to 
1937 were listed by register number. 



SYSTEMATIC DESCRIPTION OF THE 
AMMONITES AND NAUTILOIDS 

This section is not intended be a full description of the ammonites in 
the Sinemurian and Lower Pliensbachian of Robin Hood's Bay, but 
all ammonites that have been figured before are included in a list in 
systematic order, and this gives an indication of their synonymies. 
All the Robin Hood's Bay ammonites that have been described or 
figured by the following authors are included: J. Sowerby and J. de 
C. Sowerby (1812-1846), Young & Bird (1822, 1828), Phillips 
(1829, 1835, 1875), Brown (1837, 1889), Simpson (1843, 1855, 
1884), Blake (1876), Wright (1878-82), Hyatt (1889), Buckman 
(1909-30), Spath (1923*, 1924, 1925a, 1938, 1956), Trueman & 
Williams (1925), Jaworski ( 193 1 ), Howarth (1955, 1962), Dean etal 
( 1 96 1 ), Howarth & Donovan ( 1 964), Guerin-Franiatte ( 1 966), Getty 
(1973), Donovan & Forsey (1973), Schlegelmilch (1976, 1992), 
Phelps (1985), Dommergues (1987) and Dommergues & Meister 
(1992). 

All the ammonites listed are from Robin Hood's Bay. except 
where indicated otherwise, and the beds from which the type and 
figured specimens might have come are identified with varying 
degrees of confidence, as indicated in the list; register numbers are 
given, where known. The list also shows the data on which the 
identifications in the paper are based (eg. by giving references to 
the type specimens in most cases, including those that are not 
Yorkshire specimens). 56 of the better preserved ammonites in 
Bairstow's collection are figured to illustrate the identifications and 
the contents of some of the subzones. Further discussion of 
synonymies, identifications and distribution in the zones and 
subzones is found in the section on Biostratigraphy, and more 
details of the identifications of the type specimens of some species 
can be found in Howarth (1962). All measurements are in millime- 
tres (mm); D = diameter, Wh = whorl height, Wb = whorl breadth, 
U = diameter of the umbilicus. 



Order AMMONOIDEA Zittel, 1884 
Suborder PHYLLOCERATINA Arkell, 1950 
Family JURAPHYLLITIDAE Arkell, 1950 
Genus TRAGOPHYLLOCERAS Hyatt, 1900 

Tragophylloceras numismale (Quenstedt, 1845) 

PI. 1 , fig. 1 

1 843 Ammonites huntoni Simpson: 41 . 

1 845 Ammonites heterophyllus numismalis Quenstedt: 1 00, pi. 6, 

figs 4a, b, 5a, b, non figs 3a, b, 5c (figs 5a, 5b, from 

Germany, designated lectotype by Buckman, 1912: viii). 
1855 Ammonites nanus Simpson: 38. 
1921 Tragophylloceras huntoni (Simpson); Buckman: pi. 219 

(paratype or holotype, WM 477; ?from bed 5 17 or 520). 
1926 Tragophylloceras nanum (Simpson); Buckman: pi. 679 

(holotype, WM 472; from bed 517 or 520). 
1964 Tragophylloceras numismale (Quenstedt); Howarth & 

Donovan: 295, pi. 48, fig. 5 (BM C.67766; from bed 517 or 

520). 

Range. Beds 505.2-544.5, Taylori to Polymorphus Subzones; 1 7 
specimens. 

Tragophylloceras loscombi (J. Sowerby, 1817) 

1817 Ammonites loscombi J. Sowerby: 185, pi. 183. 

1843 Ammonites ambiguum Simpson: 8. 

1843 Ammonites robinsoni Simpson: 42. 

1910 Rhacoceras ambiguum (Simpson); Buckman: pi. 16 
(holotype, WM 89; ?from bed 569). 

1914 Tragophylloceras loscombi (J. Sowerby); Spath: 336, pi. 
49, fig. 1 (holotype, from Dorset). 

1921 Tragophylloceras robinsoni Buckman: pi. 220 (paratype, 
WM 478; ?from bed 569). 

1964 Tragophylloceras loscombi (J. Sowerby); Howarth & Do- 
novan: 301, pi. 49, figs 4-7 (from Dorset). 

Range. Found in bed 569 only, Masseanum Subzone; 2 speci- 
mens. 

REMARKS. This single specimen high in the Masseanum Subzone 
is at a lower horizon than specimens in Dorset, where they have not 
been recorded from below the Luridum Subzone (Howarth & Dono- 
van, 1964: 293, 302). 



Suborder LYTOCERATINA Hyatt, 1889 

Superfamily LYTOCERATACEAE Neumayr, 1875 

Family LYTOCERATIDAE Neumayr, 1875 

Genus LYTOCERAS Suess, 1865 



Lytoceras fimbriatum (J. Sowerby, 1817) 



PI. 1, fig. 3 



1817 Ammonites fimbriatus J. Sowerby: 145, pi. 164. 
1919 Fimbrilytoceras fimbriatum (J. Sowerby); Buckman: pi. 
130A-C (from Dorset). 

Range. Beds 570-578.5, Ibex Zone; 25 specimens. Two Lytoceras 
of indeterminate species were found in beds 568 (top) and 584. 

Remarks. Lytoceras fimbriatum is confined to the Ibex Zone in 
Robin Hood's Bay, except for one poorly preserved specimen in bed 
584 (Maculatum Subzone) that can only be determined as Lytoceras 
sp. indet. Many of those in the Ibex Zone are large and well- 
preserved, and one of the best specimens is figured in PI. 1. fig. 3. 
Sowerby's figured specimen, now lost, was from Dorset. 



LOWER LIAS OF ROBIN HOOD'S BAY 



119 



Suborder AMMONITINA Zittel, 1884 

Superfamily PSILOCERATACEAE Hyatt, 1867 

Family PSILOCERATIDAE Hyatt, 1867 

Blocks of limestone containing Psiloceras, Caloceras and other 
Hettangian and Lower Sinemurian ammonites are sometimes found 
loose in Robin Hood's Bay, and species have been described by 
several authors as coming from 'Robin Hood's Bay'. They are not 
from the inter-tidal exposures (the lowest of which is in the upper 
part of the Semicostatum Zone), but are derived from Glacial Drift 
nodules that are widespread in the bay and were exploited by 19th 
century collectors. 

Genus PSILOCERAS Hyatt, 1867 

Psiloceras erugatum (Phillips, 1829) 



1829 



1962 



Ammonites erugatus Phillips: 163, pi. 13, fig. 13; also 
Phillips, 1835: 135, pi. 13, fig. 13; and Phillips, 1875:270, 
pi. 13, fig. 13. 

Psiloceras erugatum (Phillips); Howarth: 99, pi. 14, fig. 2 
(holotype, BM 37982, from 'Robin Hood's Bay'). 



Psiloceras aff. sampsoni (Portlock, 1843) 

1879/81 Aegoceras planorbis (J. Sowerby); Wright: 308 (1881), pi. 
14, figs 1,2 (1879) (SMJ18216, from 'Robin Hood's Bay'). 

Genus CALOCERAS Hyatt, 1870 

Caloceras belcheri (Simpson, 1843) 

1843 Ammonites belcheri Simpson: 12. 

1910 Caloceras belcheri (Simpson); Buckman: pi. 17 (holotype, 

WM 101, from Robin Hood's Bay). 
1879/81 Aegoceras belcheri (Simpson); Wright: 313 (1881), pi. 15, 

figs 7, 8 (1879) (SMJ18217, from Robin Hood's Bay), 9. 
1976 Psiloceras (Caloceras) johnstoni (J. de C. Sowerby); 

Schlegelmilch, 1976: 106, pi. 5, fig. 8 (WM 101). 

Caloceras convolutum (Simpson, 1855) 

1855 Ammonites convolutus Simpson: 43 (non Ammonites 

convolutus Schlotheim, 1820). 
1910 Caloceras convolutum (Simpson); Buckman 

(holotype, WM 491, from Robin Hood's Bay). 



Caloceras wrighti Spath, 1924 

1880/81 Aegoceras belcheri (Simpson); Wright: 313 (1881), pi. 19, 
figs 1,2(1880) (holotype, from North Cheek, Robin Hood's 
Bay, ?lost). 

1924 Caloceras wrighti Spath: 191 (nom. nov. for Wright's fig- 
ured specimen). 



Family SCHLOTHEIMIDAE Spath, 1923 
Genus SCHLOTHEIMIA Bayle, 1878 

Schlotheimia redcarensis (Young & Bird, 1822) 



1822 



1925 



Ammonites redcarensis Young & Bird: 248, pi. 14, fig. 13; 
also Young & Bird, 1828: 258, pi. 14, fig. 10. 
Schlotheimia redcarensis (Young & Bird); Buckman, pi. 
608 (neotype, WM 314; if from Robin Hood's Bay, as 
labelled, it must be from Glacial Drift). 



Genus SAXOCERAS Lange, 1924 

Saxoceras aequale (Simpson, 1855) 

1855 Ammonites aequalis (Simpson): 49. 

1925a Saxoceras aequale (Simpson); Spath: 204, fig. 4 (drawing 
of holotype, BM 18109). 

1962 Saxoceras aequalis Howarth: 100, pi. 14, fig. 3 (holotype, 
BM 1 8 109, from Robin Hood's Bay or Redcar, more prob- 
ably the latter). 

Genus ANGULATICERAS QuemtedU 1883 

Angulaticeras sulcatum (Simpson, 1843) 

1843 Ammonites sulcatus Simpson: 55 (non Ammonites sulcata 

Lamarck, 1822). 
1911 Schlotheimia sulcata (Simpson); Buckman: pi. 38 (holotype, 

WM 743; ?from beds 461^164). 
1976 Angulaticeras sulcatum (Simpson); Schlegelmilch: 1 12, pi. 

8, fig. 7 (WM 743). 

RANGE. Six small examples of Angulaticeras sp. indet. were found 
in beds 461^-64.33 and 485.2, Denotatus to Oxynotum Subzones. 

Genus Macrogrammites Buckman, 1928 

Macrogrammites antiquatum (Simpson, 1855) 

1855 Ammonites antiquatus Simpson: 36. 

1927 Schlotheimia antiquata (Simpson); Buckman, 1927, pis 
718A, B (holotype, WM 79/80); if this holotype is a large 
smooth outer whorl of the Hettangian genus 
Macrogrammites as identified by Buckman (1928: on cap- 
tion to pi. 7 1 8A* [re-issue of 1 927, pi. 7 1 8 A]), then it comes 
from Glacial Drift in Robin Hood's Bay, or from Redcar; it 
is not from beds exposed in Robin Hood's Bay. 



Family ARIETITIDAE Hyatt, 1875 

Subfamily ARIETITINAE Hyatt, 1875 

Genus CORONICERAS Hyatt, 1867 

Subgenus ARIETITES Waagen, 1869 



pi. 18 Coroniceras (Arietites) alcinoe (Reynes, 1879) 

PL 1, fig. 8 

1879 Ammonites alcinoe Reynes: pi. 23, figs 7, 8, 9-1 1 (neotype, 

from France). 
1955 Pararnioceras alcinoe (Reynes); Donovan: 14. 

Range. Beds 424.3^429.64, Sauzeanum Subzone; 5 specimens. 

Coroniceras (Arietites) validanfractum (Simpson, 1855) 

1855 Ammonites validanfractus Simpson: 95. 

1962 Coroniceras validanfractum Howarth: 101, pi. 14, figs 4 
(holotype, WM 282), 5 (paratype, WM 530); similarly 
preserved specimens have been found at Redcar, but not at 
Robin Hood's Bay; this species is close to C. (A.) alcinoe 
(Reynes). 

Coroniceras (Arietites) obesulus (Blake, 1876) 

1876 Arietites obesulus Blake: 284, pi. 5, fig. 2 (lectotype, SM 
J34800, from Robin Hood's Bay); also very similar to C. 
(A.) alcinoe (Reynes). 



120 



PLATE 1 



M.K. HOWARTH 




LOWER LIAS OF ROBIN HOOD'S BAY 

ICoroniceras (Arietites) radiatus (Simpson, 1843) 

1 843 Ammonites radiatus Simpson: 47 (non Ammonites radiatus 

Bruguiere, 1789). 
1911 Arietites radiatus (Simpson); Buckman: pi. 35 (holotype, 

WM 304, possibly from beds 426^129, but it is only 1 2 mm 

diameter and is not identifiable). 

Coroniceras (Arietites) cf. planaries (Reynes. 1879) 

1878/81 Arietites nodulosus (Young & Bird); Wright: 288 (1881), 
pi. 6, figs 2, 3 (1878) (C.1880, possibly from beds 426- 
429). 

Genus ARNIOCERAS Hyatt, 1867 

Arnioceras semicostatum (Young & Bird, 1828) 

PI. 1, fig. 2 

1828 Ammonites semicostatus Young & Bird: 257, pi. 12, fig. 10. 

1855 Ammonites vetustus Simpson: 88. 

1876 Arietites semicostatus (Young & Bird); Blake: 288, pi. 5. 

fig. 4a (upper figure, BM C. 17935; lower figure, BM 

C. 17934 (?malformation); both from Redcar). 
1876 Arietites difformis (Emmrich); Blake: 289, pi. 6, fig. 3 

(C. 17933, possibly from Redcar). 
1889 Arnioceras semicostatum (Young & Bird); Hyatt: 165, pi. 2, 

figs 12, 13 (from 'Whitby', presumably from Robin Hood's 

Bay). 
1918 Arnioceras semicostatum Buckman: pi. 1 12 (holotype, WM 

924). 
1925a Arnioceras semicostatum (Young & Bird); Spath: 329, fig. 

10a(BMC.86a). 
1931 Arnioceras semicostatum (Young & Bird); Jaworski: 1 1 1, 

pi. 5, fig. 1 (WM 924). 
1956 Arnioceras semicostatum (Young & Bird); Spath: 153, pi. 

10, figs 6 (BM C.25651), 7 (BM C.17933). 

1961 Arnioceras semicostatum (Young & Bird); Dean et al.\ pi. 
65, fig. 4 (BM C.25651). 

1962 Arnioceras vetustum (Simpson); Howarth: 103, pi. 15, fig. 
2 (holotype, GSM 26404, ?from bed 421 or 424). 

1976 Arnioceras semicostatum (Young & Bird); Schlegelmilch: 
138, pi. 21, fig. 4 (WM 924). 

RANGE. Beds 42 1 .4-^129.64, Sauzeanum Subzone; 1 1 8 specimens. 

Remarks. Jaworski's (1931: pi. 5, fig. 1) enlarged figure of the 
holotype is much better than Buckman's (1918) figure. Older collec- 
tions contain many well-preserved Arnioceras semicostatum from 
limestone nodules. Such good specimens were not found by Bairstow 
and they have sometimes been considered to come from Glacial 



121 

Drift, but judging from the preservation it is possible that they came 
from nodules near the top of bed 42 1 .4 or from beds 424.2 or 424.3. 
Although Arnioceras persists elsewhere into the overlying Brooki 
and Birchi Subzones, all the well-preserved Robin Hood's Bay 
specimens appear to have come from these limestone nodules of the 
Sauzeanum Subzone. One of the better examples in Bairstow's 
collection is figured in PI. 1, fig. 2. 

The range of Arnioceras is extended down to bed 42 1 . 1 and up to 
beds 430-436 by six specimens of indeterminate species found by 
Bairstow. 

Arnioceras acuticarinatum (Simpson, 1855) 

1855 Ammonites acuticarinatum (Simpson): 94) 

1911 Arnioceras acuticarinatum (Simpson); Buckman: pi. 40 

(holotype, WM 295; ?from beds 421^24). 
1 93 1 Arnioceras acuticarinatum (Simpson): Jaworski: 1 26, pi. 5, 

fig. 2 (WM 295). 
1976 Arnioceras acuticarinatum (Simpson); Schlegelmilch, 

1976: 138, pi. 21, fig. 3 (WM 295). 

REMARKS. The best figure of the holotype is that of Jaworski, and 
such a well-preserved specimen might have come from beds 421 — 
424. Its more finely ribbed inner whorls possibly separate it from A. 
semicostatum. 

Arnioceras miserabile (Quenstedt, 1856) 



1856 
1876 

1884 

1925« 



1976 



Ammonites miserabilis Quenstedt: 71, pi. 8, fig. 7. 
Aegoceras nigrum Blake: 274, pi. 6, fig. 6 (lectotype, BM 
C. 17889, possibly from bed 424.2 or 424.3). 
Ammonites miserabilis Quenstedt; Quenstedt: 106, pi. 13, 
figs 27-30. 

Arnioceras nigrum (blake); Spath: 329, fig. 10b (BM 50150c, 
possibly from bed 424.2 or 424.3). 

Arnioceras miserabile (Quenstedt); Guerin-Franiatte: 254, 
pi. 136, figs 1 (neotype, from Germany, original of Quen- 
stedt, 1 884: pi. 21 , fig. 27), 2-4 (from Germany and France). 
Arnioceras miserabile (Quenstedt); Schlegelmilch: 49, pi. 
21, fig. 5 (neotype). 



RANGE. Beds 424.2^425.5, Sauzeanum Subzone; 7 specimens. 
Genus VERMICERAS Hyatt, 1889 

Vermiceras multanfractum (Simpson, 1855) 

1855 Ammonites multanfractus Simpson: 95. 

1962 Vermiceras multanfractum (Simpson); Howarth: 101, pi. 
14, fig. 6 (neotype, WM 281); not found by Bairstow, possibly 
derived from Glacial Drift in Robin Hood's Bay. 



PLATE 1 

Fig. 1 Tragophylloceras numismale (Quenstedt). Bed 505.2, CA 2744. 

Arnioceras semicostatum (Young & Bird). Bed 424.2, CA 2830. 

Lytoceras fimbriatum (J. Sowerby). Bed 576.4, CA 2778, x 0.6; the large outer whori is part of the body chamber. 

Asteroceras confasum Spath. Bed 446.32, CA 3010. 

Euagassiceras resupinatum (Simpson). Bed 422.2, CA 2948. 

Eparietites impendens (Young & Bird). Bed 461, CA 3228. 

Aegasteroceras sagittarium (Blake). Bed 456, CA 3124. 

Coroniceras (Arietites) alcinoe (Reynes). Bed 429.64, CA 2803, x 0.6; the asterisk marks the probable end of the phragmocone. 
All figures natural size, except Figs 3 and 8. 
Where determinable the position of the end of the phragmocone is marked with an asterisk (*) on all specimens figured in the plates. 



Fig. 2 
Fig. 3 
Fig. 4 
Fig. 5 
Fig. 6 
Fig. 7 
Fig. 8 



PLATE 2 



M.K. HOWARTH 




LOWER LIAS OF ROBIN HOOD'S BAY 



123 



Subfamily AGASSICERATINAE Spath. 1924 
Genus AGASSICERAS Hyatt, 1875 

Well-preserved Agassiceras scipionianum (d'Orbigny) and the 
smaller, smoother species A. personation have been figured before 
from Robin Hood's Bay and are common in old collections, but none 
were found by Bairstow. Agassiceras mainly characterizes the 
Scipionianum Subzone, just below the lowest horizon exposed in 
Robin Hood's Bay, so, although it is possible that some of them came 
from beds 421 or 422, it is more likely that they were derived from 
Glacial Drift. 

Agassiceras personatum (Simpson, 1843) 

1843 Ammonites personatus Simpson: 9. 

1920 Agassiceras personatum (Simpson); Buckman: pi. 187, figs 
1, 2 (holotype, WM 2125), 3, 4 (paratype WM 67). 

Remarks. A small smooth species. 
Agassiceras scipionianum (d'Orbigny, 1844) 



1844 
71855 

1876 



1961 



71962 



1994 



Ammonites scipionianus d'Orbigny: 207, pi. 51, figs 7, 8. 

Ammonites Hiatus Simpson: 39 (non Ammonites Hiatus 

Simpson, 1843: 10). 

Arietites scipionianus (d'Orbigny); Blake: 287, pi. 5, fig. 3 

(BM C. 17909; locality not recorded, but the preservation is 

like that of other specimens from Robin Hood's Bay). 

Agassiceras scipionianum (d'Orbigny); Dean etai: pi. 65, 

fig. 3 (BM 37909). 

Agassiceras illatum (Simpson, 1855, non 1843); Howarth: 

102, pi. 15, fig. 1 (holotype, WM 84). 

Agassiceras scipionianum (d'Orbigny); Fischer: 53, pi. 16, 

figs 1 (lectotype), 2 (both from France). 



Agassiceras decipiens (Spath, 1923) 

1923b Aetomoceras decipiens Spath: 72. 

Remarks. Spam's three syntypes are BM C. 22067a, C. 22067b 
and a specimen at the top of the block WM 67 as figured by Buckman 
(1920: pi. 187, fig. 3), all from Robin Hood's Bay. Of these, C.22067a 
is here designated lectotype; it is similar to A. scipionianum 
(d'Orbigny), but has slightly more ribs. 

Genus EUAGASSICERAS Spath, 1924 



Euagassiceras resupinatum (Simpson, 1843) 



PL 1, fig. 5 



1843 Ammonites resupinatus Simpson: 15. 

1844 Ammonites sauzeanus d'Orbigny: 304, pi. 95, figs 4, 5. 
1855 Ammonites transformatus Simpson: 91. 

1889 Coroniceras sauzeanum (d'Orbigny); Hyatt: 184, pi. 6, figs 
4,6-9. 



1909 Agassiceras resupinatum (Simpson); Buckman: pi. 6 
(holotype, WM 96; ?from bed 421 or 422). 

1913 Agassiceras transformatum (Simpson); Buckman: pi. 75 
(holotype. WM 279, ?from bed 421 or 422). 

1913 Defossiceras defossum (Simpson, 1843); Buckman: pi. 76 
( WM 103, a paralectotype, presumably also from beds 421 — 
422; see Donovan & Forsey (1973: 13) for interpretation of 
Defossiceras and its type species). 

1994 Euagassiceras sauzeanum d'Orbigny; Fischer: 84, pi. 16, 
fig. 3 (holotype, from France). 

Range. Beds 421.4-426.1, Sauzeanum Subzone; 38 specimens, 
of which the best preserved are in beds 42 1 and 422. 

REMARKS. Typical examples of Euagassiceras occur in beds 421 
and 422, and it is likely that the holotypes of Simpson's species E. 
resupinatum and E. transformatum are from those beds; both species 
are synonyms off. sauzeanum (d'Orbigny, 1844), and E. resupinatum 
(Simpson, 1843) is the senior name for this species. 



Subfamily ASTEROCERATINAE Spath 1946 
Genus ASTEROCERAS Hyatt, 1867 



Asteroceras obtusum (J. Sowerby, 1817) 

1817 Ammonites obtusus J. Sowerby: 151, pi. 167. 

1966 Asteroceras obtusum (J. Sowerby); Guerin-Franiatte: 294, 

pi. 170 (lectotype, Oxford University Museum, OUM 

J. 1 194, from Charmouth, Dorset). 

RANGE. One specimen in bed 446.32, Obtusum Subzone. A single 
Asteroceras sp. indet. in bed 446.3 1 is the lowest recorded Asteroceras 
in Robin Hood's Bay. 



Asteroceras confusum Spath, 1925 



PI. 1, fig. 4; PI. 2, fig. 1 



I Arietites obtusus (J. Sowerby); Wright: 293 (1881), pi. 21, 
figs 3,4(1 880) (holotype of Asteroceras confusum Spath). 

1925rt Asteroceras confusum Spath: 300. 

1966 Asteroceras confusum Spath; Guerin-Franiatte: 296, pi. 172 
(holotype, BM C.2223, from Bredon, Worcestershire). 

Range. Five specimens in beds 446.32 and 446.33, Obtusum 
Subzone. 

Remarks. A. confusum has thicker whorls and more prominent 
grooves bordering the keel on the venter than A. obtusum. 



Asteroceras stellare (J. Sowerby, 1815) 

1815 Ammonites stellaris J. Sowerby: 2 1 1 , pi. 93. 

1880/81 Arietites stellaris (J. Sowerby); Wright: 295 (1881), pi. 22, 

figs 3-5 (1880) (lectotype, BM 43969a, from Dorset). 
1882 Aegoceras sagittarium (Blake); Wright: 355, pi. 52, figs 1- 



PLATE 2 

Fig. 1 Asteroceras confusum Spath. Bed 446.32, CA 301 1 , x 0.6. 

Fig. 2 Asteroceras blakei Spath. Bed 452, CA 2990, x 0.5. 

Figs 3, 5 Caenisites turneri (J. de C. Sowerby). Bed 433.3. 3, CA 3194. 5, CA 3197; the asterisk marks the probable end of the phragmocone. 

Fig. 4 Aegasteroceras crassum Spath. Bed 458.2, CA 3048; a body chamber fragment. 

Fig. 6 Gagaticeras neglectum (Simpson). Bed 468, CA 3326; probably wholly septate. 

Fig. 7 Gagaticeras exortum (Simpson). Bed 468, CA 3292; the asterisk marks the probable end of the phragmocone. 

Fig. 8 Eparietites bairstowi sp. nov. Paratype, bed 455.5, CA 3218; probably wholly septate. 

All figures natural size, except Figs 1 and 2. 



124 



M.K. HOWARTH 



1961 



3 (BM C.3124, a large specimen that is very similar to 
specimens from bed 45 1 ; it is more involute and has higher 
whorls than sagittahum). 

Asteroceras stellare (J. Sowerby); Dean etal. : pi. 67, fig. 2 
(lectotype, BM 43969a, from Dorset). 



Range. Beds 447-45 1, Stellare Subzone; 19 specimens. 

Remarks. A. stellare is a large species, and is more involute and 
has more massive whorls than A. obtusum. 



Asteroceras blakei Spath, 1925 

1925a 



PI. 2, fig. 



Asteroceras blakei Spath: 264, fig. 5 (holotype, BM C. 1999 1 ; 
from beds 452^155.4, probably 452-53). 
1925a Asteroceras marstonense Spath: 267, fig. 7 (holotype, BM 
37948, probably from bed 452 or 453). 

Range. Beds 452-455.4, Stellare and Denotatus Subzones; 29 
specimens. 

Remarks. A. blakei occurs at a higher stratigraphical level, and is 
more involute and more compressed than A. obtusum. A. marstonense 
is a synonym, and its holotype comes from Robin Hood's Bay 
despite being named after Marston Magna, Somerset. 

Genus AEGASTEROCERAS Spath. 1925 



Aegasteroceras sagittarium (Blake, 1876) 



1876 



PL 1. fie. 7 



Aegoceras sagittarium Blake: 276, pi. 7, figs 2A (lectotype. 

SM J 18230), 2B (paralectotype, BM C. 1 788 1 ). 
1880/82 Aegoceras acuticostatum Wright: 371 (1882), pi. 35,figs 1- 

3 (1880) (holotype, SM J 18230); objective synonym of 

Aegoceras sagittarium Blake, 1876. 
1882 Aegoceras sagittarium Blake; Wright: 355, pi. 52, figs 4, 5 

(BM C.1873); pi. 52A, figs 3, 4 (BM C.1873), 5 (BM 

C.1874), 6 (BMC. 1875). 
1925a Aegasteroceras simile Spath: 265, fig. 6a (holotype, BM 

C.26687). 
1966 Aega.?ferocera.s , «7M//eSpath;Guerin-Franiatte:310,pl. 189 

(BM C.26687). 
1966 Aegasteroceras sagittarium (Blake); Guerin-Franiatte: 312, 

pi. 192, fig. 1 (SMJ18230). 

Range. Beds 454. 1-458.1, Stellare and Denotatus Subzones; 123 
specimens; all the figured ones are from beds 454.2 or 456, probably 
the latter. 



Aegasteroceras crassum Spath, 1925 



PL 2, fig. 4 



1 882 Aegoceras sagittarium Blake; Wright: 355, pi. 52A, figs 1 , 
2 (BM C.1922, from bed 458.2 or 458.3, holotype of 
Aegasteroceras crassum Spath, 1925). 

1925a Aegasteroceras crassum Spath: 266. 

Range. Beds 449^158.3, Stellare and Denotatus Subzones; 9 
specimens. 

Genus CAENISITES Buckman, 1925 

Caenisites turneri (J. de C. Sowerby, 1824) PL 2, figs 3, 5 

1824 Ammonites turneri J. de C. Sowerby: 75, pi. 452, upper 

figure. 
1879/81 Arietites turneri (J. de C. Sowerby): Wright: 292 (1881), pi. 

12, fig. 4 ( 1 879) (lectotype, BM 43973a, from Glacial Drift, 

Norfolk). 



1956 



1961 



Euasteroceras turneri (J. de C. Sowerby); Arkell: 760. pi. 
31, fig. 1 (BM 43973a). 

Caenisites turneri (J. de C. Sowerby); Dean et al.\ pi. 66, 
fig. 2 (from Bredon Hill. Worcestershire). 



Range. From beds 433.3 and 435, Birchi Subzone: 26 specimens. 

Caenisites brooki (J. Sowerby, 1818) 

1818 Ammonites brooki J. Sowerby: 203. pi. 190. 

1961 Caenisites brooki (J. Sowerby); Dean et al.\ pi. 66, fig. 1 

(BM C.5606, from Charmouth, Dorset). 
1966 Caenisites brooki (J. Sowerby); Guerin-Franiatte: 325, pi. 

210 (holotype, OUM J. 16020, from Lyme Regis, Dorset). 

Range. From beds 431.2^133.3, Brooki and Birchi Subzones; 1 1 
specimens. 

Remarks. One of those from bed 431.3 is notable in being 
extremely large: before removal from the rock, it was measured by 
Bairstow as 585 mm diameter. The outer part was not recovered, but 
the portion now preserved in the collection is 445 mm diameter and 
has a massive, almost smooth outer half whorl with keel and grooves 
on the venter. The inner whorls are covered in matrix and probably 
crushed. 

Genus EPARIETITES Spath, 1924 

Eparietites impendens (Young & Bird, 1828) 

PI. 1, fig. 6; PL 4, fig. 1 

1828 Ammonites impendens Young & Bird: 266. 

1855 Ammonites denotatus Simpson: 76. 

1855 Ammonites tenellus Simpson: 97. 

1876 Arietites impendens (Young & Bird); Blake: 290, pi. 6, fig. 

7 (BMC. 17936). 
1878/81 Arietites collenoti (d'Orbigny); Wright: 304 (1881), pi. 6 

(1878), fig. 1 (BMC. 1881; probably from bed 464.32), pi. 

22B, figs 1-3 (holotype of Ammonites denotatus, SM J3273). 
1912 Arietites tenellus (Simpson); Buckman: pi. 54 (holotype, 

WM 293, probably from bed 461 or 464.32). 
1912 Arietites denotatus (Simpson); Buckman: pis 67A, B 

(holotype, SM J3273). 
1919 Arietites impendens (Young & Bird); Buckman: pi. 120 

(holotype, WM 292, from bed 462 or 464. 1 ). 
1961 Eparietites denotatus (Simpson); Dean et ah: pi. 66, fig. 4 

(BMC.17936). 

Range. Beds 457-464.32, Denotatus and Simpsoni Subzones; 57 
specimens; all the figured specimens are from beds 461^64. 

Remarks. Both denotatus and tenellus are synonyms of E. 
impendens; the best type specimen is the holotype of Simpson's 
denotatus. but it is clearly the same as the holotype of Young & Bird's 
impendens. and the holotype of tenellus differs only in having the 
crushed-flat type of preservation that is common at some horizons. 



Eparietites bairstowi sp. nov. 



PL 2, fig. 8; PL 3 



Holotype. CA 3217, from bed 455.2, lower part of Denotatus 
Subzone. 

Paratypes. CA 3218 and 3219, from bed 455.5, lower part of 
Denotatus Subzone. 

DIAGNOSIS. An evolute species of Eparietites in which the umbili- 



LOWER LIAS OF ROBIN HOOD'S BAY 



125 



cal width is 38^40% of the diameter; the whorls are quickly expand- 
ing and massive, the whorl breadth is large, and the venter is 
tricarinate-bisulcate, but loses the sulci at the largest sizes; inner 
whorls have radial ribs and small ventro-lateral tubercles, but all 
ornament fades by 230 mm diameter. 

Description. Before removal from the rock, Bairstow measured 
the holotype as having a crushed outer whorl ending at 480 mm ('19 
inches' ) diameter. At that size it was probably complete, but the part he 
collected in 1933 is solid and uncrushed up to the end of the 
phragmocone at 330 mm diameter, followed by a short portion at the 
begining of the crushed body-chamber ending at about 350 mm 
diameter. Slightly more than one whorl up to the end of the phragmocone 
is preserved, to which is attached a small portion of the upper part of the 
side of the next inner whorl. These whorls are massive, rapidly expand- 
ing, moderately evolute, and have a whorl section in which the whorl 
sides converge slightly towards the tricarinate-bisulcate venter, which 
has a strong central keel. Moderately strong radial ribs fade and 
disappear three-quarters of a whorl before the end of the phragmocone, 
and the remaining whorls are smooth. 

The larger paratype (CA 3218) consists of inner whorls up to 40 
mm diameter, but only a quarter of a whorl up to 25 mm is well 
preserved. This has quadrate, moderately evolute whorls with a 
tricarinate-bisulcate venter, and strong straight radial ribs up to small 
ventro-lateral tubercles; the ribs then bend strongly forwards on the 
side of the venter and join the lateral keels. 

The smaller paratype (CA 3219) consists of small inner whorls up 
to only 12.5 mm diameter. Its whorl shape, ribs and tubercles are 
similar to those of the larger paratype. 

Measurements (in mm) 





D 


Wh 


Wb 


U 


CA3217 


342 


116(0.34) 


90 (0.26) 


136(0.40) 


CA3217 


265 


92 (0.35) 


67 (0.25) 


101(0.38) 



Remarks. This is the oldest Eparietites and is more evolute than 
any of the succeeding species. The large holotype has massive 
whorls, with a quadrate whorl section in which the whorl sides 
converge only slightly towards the venter, a large whorl breadth, and 
a tricarinate-bisulcate venter; the sulci bordering the central keel on 
the venter slowly disappear at the largest sizes. E. impendens occurs 
higher up at Robin Hood's Bay and has much more compressed and 
involute whorls, the umbilical width being 22-24% of the diameter 
compared with 38^4-0% in E. bairstowi. 

Many Eparietites occur in the Denotatus Subzone in the top 0.5 m 
of the Frodingham Ironstone at Scunthorpe, Lincolnshire. Speci- 
mens attain very large sizes, some in the NHM collections being up 
to 600 mm diameter. Most of the smaller specimens are E. impendens, 
but the larger specimens belong mainly to the more evolute species 
Eparietites undaries (Quenstedt), in which the umbilical width is 
27-34% ofthe diameter (Guerin-Franiatte, 1966: 3 19). There is also 
one good example off. bairstowi in the Frodingham Ironstone: it is 
a quarter whorl uncrushed fragment, wholly septate, with a whorl 
height and breadth of 103.5 and 69.3 mm respectively, and is closely 
similar to the Robin Hood's Bay holotype at the same size. 

Genus EPOPHIOCERAS Spath, 1924 

Epophioceras landrioti (d'Orbigny, 1849) 

1849 Ammonites landrioti d'Orbigny: 213. 
1907 Ammonites landrioti d'Orbigny; Thevenin: 94, pi. 7, figs 4, 
5 (holotype, from the Obtusum Zone, France). 



1966 Epophioceras landrioti (d'Orbigny); Guerin-Franiatte: 329, 
pi. 217 (holotype). 

Range. Beds 446.5-448.11, Obtusum and Stellare Subzones; 3 
specimens. A single Epophioceras sp. indet. was found in bed 446.4. 



Family ECHIOCERATIDAE Buckman, 1913 
Genus PALAEOECHIOCERAS Spath, 1929 



Palaeoechioceras sp. indet. 

1973 Palaeoechioceras sp.; Getty: 9, pi. 1, figs 1 (BM C.79680), 
5 (BM C.79678), 9 (BM C.79679); all from bed 467. 

RANGE. Bed 467, Simpsoni Subzone; 3 specimens. 

Genus GAGAT1CERAS Buckman, 1913 

REMARKS. In Robin Hood's Bay examples of Gagaticeras are 
found only in beds 467^170, the top half of the Simpsoni Subzone. 
Although they are divided here into G. neglectum with medium to 
coarse ribs, G. finitimum with finer ribs, G. exortum with strongly 
rursiradiate ribs, and G. gagateum with markedly depressed whorls, 
larger collections of better specimens might suggest that there are 
fewer than four species present. 

Gagaticeras gagateum (Young & Bird, 1828) 

1828 Ammonites gagateus Young & Bird: 255, pi. 12, fig. 7. 
1876 Aegoceras gagateum (Young & Bird); Blake: 275, pi. 6, fig. 

8 (BM C. 17883, from bed 467). 
1 880/82 Aegoceras gagateum (Young & Bird); Wright: 364, pi. 37, 

figs 8, 9 (BM C.2228, probably from bed 467). 
1913 Gagaticeras gagateum (Young & Bird); Buckman: pi. 78 

(holotype, WM 104, from bed 467). 
1919 Gagaticeras funiculatum Buckman: pi. 122 (holotype, BM 

C.41783). 
1962 Gagaticeras gagateum (Young & Bird); Howarth: 102, pi. 

14, fig. 6 (WM 744, paratype of Ammonites multanfractus 

Simpson, 1855). 
1976 Gagaticeras gagateum (Young & Bird); Schlegelmilch: 

138, pi. 21, fig. 7 (WM 104). 

RANGE. Occurs only in bed 467, Simpsoni Subzone; 2 specimens; 
has strongly depressed whorls. 

Gagaticeras exortum (Simpson, 1855) PI. 2, fig. 7 

1855 Ammonites exortus Simpson: 44. 

1855 Ammonites integricostatus Simpson: 46. 

1910 Echioceras exortum (Simpson); Buckman: pi. 19, figs 2, 3 

(neotype (designated Howarth, 1962: 106), WM 645). 
1912 Androgynoceras integricostatum (Simpson); Buckman: pi. 

47 (holotype, WM 92). 

RANGE. Beds 467-70, Simpsoni Subzone; 15 specimens; charac- 
terized by markedly rursiradiate ribbing. 



Gagaticeras neglectum (Simpson, 1855) 



PI. 2 fig. 6 



1855 Ammonites neglectus Simpson: 45. 

1914 Parechioceras neglectum (Simpson); Buckman: pi. 101 

(holotype, WM 98). 
1976 Gagaticeras neglectum (Simpson); Schleglemilch: 138, pi. 

21, fig. 8 (WM 98). 



126 

Range. Beds 467-70, Simpsoni Subzone; 77 specimens; the 
holotype came from bed 468 or 470; has medium to coarse ribbing. 

Gagaticeras finitimum (Blake, 1876) 

1876 Aegoceras (?) finitimum Blake: 273, pi. 6, fig. 9. 
1914 Parechioceras finitimum (Blake); Buckman: pi. 100A 
(holotype, SM J3280, possibly from bed 468). 

Range. Beds 467 and 468, Simpsoni Subzone; 11 specimens; 
finely ribbed. 



M.K. HOWARTH 
Genus ECHIOCERAS Bayle, 1878 



Echioceras raricostatum (Zieten, 1831) 

1831 Ammonites raricostatus Zieten: 18, pi. 13, fig. 4. 

1855 Ammonites cereus Simpson, 1855:47. 

1912 Echioceras cereum (Simpson); Buckman: pi. 49 (holotype, 

WM461). 
1973 Echioceras raricostatum (Zieten); Getty: 13, pi. 1, fig. 7 

(neotype, designated Getty, from Wurttemberg, Germany). 



!* 





• . 



- 
- 



& 



!£^- 






m^ri 



ijr 









88 s m ' P - 



PLATE 3 

Eparietites bairstowi sp. nov. Holotype, bed 455.2. CA 3217, x 0.48. 



LOWER LIAS OF ROBIN HOOD'S BAY 



127 



Remarks. Examples of Echioceras with strongly depressed whorls 
and coarse ribs were not found by Bairstow, and the horizon of 
Simpson's holotype WM 461 is not known; presumably it ought to 
have come from the Raricostatoides Subzone. 

Echioceras raricostatoides (Vadasz, 1908) PI. 4, fig. 2 

1908 Arietites raricostatoides Vadasz: 373. 

1925 Echioceras fill gidum Trueman & Williams: 717, pi. 1, fig. 

12 (BM C. 17424, possibly from bed 489). 
1973 Echioceras raricostatoides (Vadasz); Getty: 13, pi. 1, fig. 

12 (neotype, designated by Getty, from Nancy, France). 

Range. Beds 488^19 1.1, Raricostatoides Subzone; 1 5 specimens. 

Remarks. The very well preserved specimen from bed 489 fig- 
ured in PI. 4, fig. 2 has a closely similar rib-density to that of the 
neotype shown in Getty's (1973: 15, fig. 3) graph. The more densely 
ribbed species E. aeneum Trueman & Williams, Echioceras 
laevidomum (Quenstedt; Schleglemilch, 1976: pi. 21, fig. 12, 
lectotype) which has modified ribbing at large whorl sizes, and E. 
pauli (Dumortier, 1867: pi. 29, figs 5, 6), all said to occur in the 
lowest part of the Raricostatoides Subzone by Getty (1973: 14), were 
not identified amongst Bairstow's material. 

Echioceras intermedium (Trueman & Williams, 1925) 



1925 
1973 

Range. 
mens. 



Pleurechioceras intermedium Trueman & Williams: 720, 
pi. 2 fig. 2 (holotype, BM C.26787, from bed 493. 
Echioceras intermedium (Trueman & Williams); Getty: 16, 
pi. 3, fig. 1 (BM C.79684, from bed 493). 

Beds 491.2 and 493.2, Raricostatoides Subzone; 7 speci- 



Genus LEPTECHIOCERAS, Buckman 1923 



Leptechioceras macdonnelli (Portlock, 1843) 

1843 Ammonites macdonnelli Portlock: 134, pi. 29A, fig. 12. 
1880/81 Arietites nodotianus (d'Orbigny); Wright: 301 (1881), pi. 

37, figs 3, 4 (1880) (holotype of Ammonites macdonnelli 

Portlock, from Larne, northern Ireland, Ulster Museum no. 

K8117). 
1923 Leptechioceras macdonnelli (Portlock); Buckman: pi. 443 

(BM C.41756, from Cheltenham). 
1961 Leptechioceras macdonnelli (Portlock); Dean ef al: pi. 67, 

fig. 6 (BM C.41756, from ?Larne, Co Antrim, northern 

Ireland). 
1973 Leptechioceras macdonnelli (Portlock); Getty: 16. 

Range. Beds 494^195.7, Macdonnelli Subzone; 5 specimens. 

Remarks. The earliest examples in bed 494 are identified as L. aff. 
macdonnelli because reduced ribbing persists onto their outer whorls, 
which do not become as smooth as in the later specimens from beds 
495. 13 and 495.7. Nevertheless, in those early specimens the ribs are 
more reduced than in L. nodotianum (d'Orbigny, 1843; Fischer, 
1994: 48, pi. 20, fig. 4, holotype), L. charpentieri (Schafhautl, 1 847; 
Getty, 1973: pi. 2, fig. 6, lectotype) or L. meigeni (Hug, 1 899), all of 
which are more strongly ribbed throughout. 

Genus PALTECHIOCERAS Buckman, 1924 

Paltechioceras planum (Trueman & Williams, 1925) 

1925 Leptechioceras planum Trueman & Williams: 731, pi. 2, 
fig. 5 (holotype, from Radstock, Somerset). 



1926 Leptechioceras planum Trueman & Williams; Buckman: 
pi. 696 (holotype refigured). 

Range. Beds 493.3^493.5, Raricostatoides Subzone; 5 specimens. 

Paltechioceras tardecrescens (Hauer, 1 856) 

PI. 4, figs 3, 6 

71855 Ammonites aureolus Simpson: 94. 

1856 Ammonites tardecrescens Hauer: 20, pi. 3, figs 10-12. 

1876 Arietites tardecrescens (Hauer); Blake: 285, pi. 5, figs 5 

(?BM C. 17879, from bed 498), 5b (7BM C. 17898). 
1889 Caloceras aplanatum Hyatt: 146, figs 23, 24 (on p. 147). 
1889 Arnioceras tardecrescens (Hauer); Hyatt: 168, pi. 2, fig. 19. 
71914 Echioceras aureolum (Simpson); Buckman: pi. 96 

(lectotype, designated by Donovan ( 1 958: 24), GSM 26402, 

from bed 497); ?senior synonym of P. tardecrescens. 
1926 Metechioceras aplanatum (Hyatt); Buckman: pi. 640 

(holotype, MCZ 80 (Museum of Comparative Zoology, 

Cambridge, Massachussets), from bed 498). 
1961 Paltechioceras aplanatum (Hyatt); Dean et al.: pi. 68, fig. 2 

(BM 37999, from bed 498). 
1973 Paltechioceras aplanatum (Hyatt); Getty: 21, pi. 4, fig. 1 

(BM C. 17898, from bed 498). 
1973 Paltechioceras tardecrescens (Hauer); Getty: 21, pi. 4, fig. 

2 (lectotype, designated Getty, from Adneth, Saltzburg, 

Austria). 
1992 Paltechioceras tardecrescens (Hauer); Dommergues & 

Meister: 221, figs 5(l)-5(4) (from bed 497). 

Range. Beds 497-499, Aplanatum Subzone; 228 specimens. 

Remarks. P. tardecrescens is abundant in beds 497 and 498; those 
in bed 498 are up to 175 mm diameter and are preserved in limestone 
nodules (PI. 4, fig. 6), and many of the previously figured specimens 
undoubedly came from this bed (including the holotype of Hyatt's 
species aplanatum). Most of the specimens in bed 497 are much 
smaller pyritized whorls up to about 40 mm diameter (PI. 4, fig. 3), 
though there are a few crushed and partly pyritized fragments of 
larger whorls up to 90 mm diameter. The lectotype of P. aureolum 
(Simpson, 1855; Buckman, 1914: pi. 96) is pyritized like most 
specimens in bed 497 and can be matched closely with several of 
them (eg. CA 3456 and 3480); it is only 25 mm diameter, but 
Ammonites aureolus Simpson, 1855, might be a senior synonym of 
Paltechioceras tardecrescens (Hauer, 1856). In addition, two speci- 
mens were found at the base of bed 499 and 0.08 m above the base of 
that bed respectively. 

Paltechioceras regustatum Buckman, 1914 



1911 



1914 
1973 



Echioceras aureolum (Simpson); Buckman: pi. 28 
(paralectotype of Ammonites aureolus Simpson, WM 872, 
from bed 497). 

Paltechioceras regustatum Buckman: 96c. 
Paltechioceras aureolum (Simpson); Getty: 20, pi. 5, figs 3 
(holotype of Echioceras regustatum Buckman, GSM 26439, 
from bed 496 or 497), 4 (BM C.79681, from bed 497). 



RANGE. Beds 496 and 497, Aplanatum Subzone; 18 specimens. 

Remarks. P. regustatum is the second and less common species of 
Paltechioceras in the Aplanatum Subzone, and is represented by a few 
poorly preserved specimens in bed 496 and two in bed 497; it has much 
more widely spaced ribs than P. tardecrescens at diameters of more 
than 25 mm. Two larger examples figured by Getty (1973: pi. 5, figs 3, 
4) as P. aureolum are 50 and 64 mm diameter respectively, and the latter 



128 



PLATE 4 



M.K. HOWARTH 




LOWER LIAS OF ROBIN HOOD'S BAY 



129 



was said to be from bed 497 (Getty. 1973:20. 'Tate & Blake's Jamesoni 
Zone bed 60' ). The former of Getty's specimens is the holotype of P. 
regustatum (Buckman. 1914). and this seems to be the correct specific 
name for the less densely ribbed Paltechioceras in beds 496 and 497, 
because the lectotype of P. aureolum has the same rib density as in P. 
tardecrescens, of which it might be a senior synonym. 



Family OXYNOTICERATIDAE Hyatt, 1867 
Genus OXYNOTICERAS Hyatt 1867 

REMARKS. Determination of species of Oxynoticeras in the 
Simpsoni Subzone presents many problems: Simpson ( 1843, 1855) 
proposed the specific names simpsoni, limatus, bucki, flavus and 
lens, and Spath (1925a) proposed the name eboracense, all for 
specimens from beds 467 or 468 (the type specimens of aliaenum 
and dejectum, both of Simpson, 1 855, are lost and the names are not 
usable). The name simpsoni has date and page priority, and is 
frequently used for these ammonites. However. Spath's eboracense 
also has a well-preserved type specimen, and if this represents a 
species different from O. simpsoni, then it differs only by its more 
compressed whorls up to 40 mm diameter. But the many examples of 
Oxynoticeras in beds 467 and 468 show a large amount of variation 
in whorl compression and rib strength on whorls up to 50 mm 
diameter, and there are no larger ammonites that can be identified as 
0. 'eboracense ' in having more compressed whorls than O. simpsoni 
at large sizes. O. bucki (Simpson, 1 843) and O. lens (Simpson, 1855) 
are clearly the same as O. eboracense, of which they are senior 
synonyms; O.flavum and O. limatum. both of Simpson, 1843, are 
smaller and have slightly thicker whorls like those of O. simpsoni. 
The lectotype of O. collenoti (d'Orbigny. 1844; figured by Fischer, 
1994: 85, pi. 17, fig. 3) is also very similar to the types of lens and 
eboracense . 

Larger collections of better preserved specimens will be needed to 
determine whether these Oxynoticeras can really be divided into 
more than one species, so in this paper all the specimens in beds 463- 
472.1 are identified as O. simpsoni. 

Oxynoticeras simpsoni (Simpson, 1843) PI. 4, figs 5, 8 



1843 

1843 

1843 

1843 

71855 

1876 



Ammonites simpsoni Simpson: 37. 

Ammonites limatus Simpson: 41. 

Ammonites bucki Simpson: 42. 

Ammonites flavus Simpson: 43. 

Ammonites lens Simpson: 80. 

Amaltheus simpsoni (Simpson); Blake: 291, pi. 8, fig. 4 

(BMC. 17903). 
1881/82 Amaltheus simpsoni (Bean); Wright: 392 (1882), pi. 47 

(1881), figs 4, 5 (SM J18231), 6, 7 (SM J18232). 
1912 Oxynoticeras flavum (Simpson); Buckman: pi. 55 (holotype, 

WM481). 



1912 Oxynoticeras limatum (Simpson); Buckman: pi. 56, fig. 1 

(holotype. WM 480). 
1912 Aetomoceras simpsoni (Simpson); Buckman: pis 66A, B 

(holotype, WM 813). 
1920 Oxynoticeras bucki (Simpson); Buckman: pi. 165A 

(holotype, WM 479a). 
1925« Oxynoticeras eboracense Spath: 108. 1 10. figs d,e( holotype, 

BMC. 18060). 
1925« Oxynoticeras simpsoni (Simpson); Spath: 110, figs f, g 

(BM 37998). 
1961 Oxynoticeras simpsoni (Simpson); Deanef al. : pi. 67, fig. 4 

(BMC.17903). 
71962 Gleviceras lens (Simpson): Howarth: 105, pi. 15. fig. 3 

(holotype, GSM 26405). 
1976 Oxynoticeras bucki (Simpson); Schlegelmilch: 140, pi. 22, 

fig. 12 (WM 479a). 

RANGE. Beds 463^170, Simpsoni Subzone; 63 specimens. 

REMARKS. All the figured specimens listed in the synonymy above 
probably came from beds 467 or 468. O. simpsoni is a distinctive 
species that has a larger umbilicus and thicker whorls than O. 
oxynotum and similar species. In Robin Hood's Bay it overlaps in 
stratigraphical range with Eparietites impendens, from which it 
differs mainly in whorl section: E. impendens has a differentiated 
ventral keel, flanked by narrow flat areas then angled ventro-lateral 
shoulders, and a vertical umbilical wall and rounded umbilical edge; 
in O. simpsoni the venter is either lanceolate or fastigate with no 
angles at the umbilical shoulders and without a differentiated keel, 
and the broad umbilical wall typically slopes at a low angle and 
merges gradually into the side of the whorl. E. impendens has ribs at 
least on the inner whorls; most O. simpsoni are smooth, though some 
early examples retain ribs on small inner whorls. 

The lowest O. simpsoni with no ventro-lateral angles at the side of 
the keel occurs in bed 463, where there are two large examples: one 
is part of a solid body-chamber ending at about 340 mm diameter; the 
other is 380 mm diameter and has half a whorl of body-chamber, but 
is crushed and less well-preserved. A large fragment from bed 
464.33 is septate up to at least 256 mm diameter. A smaller O. 
simpsoni from bed 464.32 is figured in PI. 4, fig. 8, which has ribbing 
on its inner whorl at about 80 mm diameter, and a small specimen 
from bed 468 is also figured (PI. 4, fig. 5). 

Oxynoticeras oxynotum (Quenstedt, 1843) PI. 4, fig. 4 

1843 Ammonites oxynotus Quenstedt: 161. 

1843 Ammonites polyophyllus Simpson: 39. 

1845 Ammonites oxynotus Quenstedt; Quenstedt: 98, pi. 5, fig. 11 

(holotype). 
1 884 Ammonites oxynotus Quenstedt; Quenstedt: 1 75, pi. 22, fig. 

29 (holotype). 
1909 Oxynoticeras podophyllum (Simpson); Buckman: pi. 8 

(holotype, WM 739). 



PLATE 4 

Fig. 1 Eparietites impendens (Young & Bird). Bed 462, CA 3243. 

Fig. 2 Echioceras raricostatoides Vadasz. Bed 489. CA 3393; wholly septate. 

Figs 3, 6 Paltechioceras tardecrescens (Hauer). 3, bed 497. CA 3572. 6, bed 498. CA 3616; probably wholly septate. 

Fig. 4 Oxynoticeras oxynotum (Quenstedt). Bed 481, CA 3716; wholly septate. 

Figs 5, 8 Oxynoticeras simpsoni (Simpson). 5, bed 468, CA 3692: wholly septate. 8. bed 464.32. CA 3652. x 0.8; the outer whorl is part of the body 

chamber. 
Fig. 7 Gleviceras doris (Reynes). Bed 476, CA 3726, x 0.6. 
All figures natural size, except Figs 7 and 8. 



130 
1961 



Oxynoticeras oxynotum (Quenstedt); Dean et al.\ pi. 66, 
fig. 5 (holotype, Geol.-Pal. Institut, Tubingen, from 
Wurttemberg, Germany). 



Range. Beds 472. 1^48 1 , Oxynotum Subzone; 4 specimens. 

Remarks. The lowest Oxynoticeras that are more involute and 
flat-whorled than O. simpsoni occur in bed 472. 1 , which is therefore 
the base of the Oxynotum Subzone. Better preserved O. oxynotum 
occur higher up in beds 475.3 and 481 (PI. 4 fig. 4), and there are 
several fragments of Oxynoticeras sp. indet. from beds 480. 482 and 
483 consisting of large compressed whorls up to 260 mm diameter 
that have acute venters and complex suture-lines. 

Other species of Oynodceras: 

lOxynoticeras aliaenum (Simpson, 1855: 85). 
lOxynoticeras dejectum (Simpson, 1855: 85). 

The type specimens of both Simpson's species are lost, and the 
species are not identifiable. 

Genus GLEVICERAS Buckman, 1918 

Gleviceras doris (Reynes, 1879) PI. 4, fig. 7 

1879 Ammonites doris Reynes: pi. 41, figs 13-15 (probably from 

France). 
1914 Oxynoticeras doris (Reynes); Pia: 7, 30, pi. 1, fig. 1; pi. 8, 

fig.l. 

Range. Beds 476 and 485.2, Oxynotum Subzone; 3 specimens. 
Gleviceras guibalianum (d'Orbigny, 1844) 



1844 
1973 



1994 



Ammonites guibalianus d'Orbigny: 259, pi. 73, figs 1—4. 

Gleviceras subguibalianum (von Pia) (sic); Donovan & 

Forsey: 9, pi. 2, fig. 1 (lectotype of Ammonites guibalianus 

d'Orbigny, designated by Donovan & Forsey, from Nantua, 

France). 

Gleviceras guibalianum (d'Orbigny); Fischer: 66, pi. 17, 

fig. 2 (lectotype refigured). 



Range. Beds 484.1^499, Oxynotum to Aplanatum Subzones; 14 
specimens. 

Remarks. Most specimens are large body-chambers, or frag- 
ments thereof, up to 300 mm diameter. 



M.K. HOWARTH 
Genus PARACYMB1TES Trueman & Williams, 1927 

Paracymbites dennyi (Simpson, 1843) 

1843 Ammonites dennyi Simpson: 9. 
1843 Ammonites arctus Simpson: 10. 
1909 Oxynoticeras dennyi (Simpson); Buckman: pi. 7, figs 1 

(lectotype, WM 470), 2, 3 (two paralectotypes). 
1911 Oxynoticeras arctum (Simpson); Buckman: pi. 36 (holotype, 

WM471). 
1966 Paracymbites dennyi (Simpson); Donovan: 3 15, pi. 53, figs 

5-12 (from Oxfordshire and Gloucestershire). 

Remarks. As revised by Donovan (1966), the holotype of this 
species should have come from the lower part of the Raricostatum 
Zone in Robin Hood's Bay, but no examples were found by Bairstow. 

Genus PAROXYNOTICERAS von Pia, 1914 

Paroxynoticeras salisburgense (Hauer, 1856) 

1856 Ammonites salisburgensis Hauer: 47, pi. 13, figs 1-3 
(lectotype, designated Donovan & Forsey (1973: 9), from 
Adneth. Austria). 

1914 Paroxynoticeras salisburgense (Hauer); Pia: 18. 73, pi. 1, 
fig. 2; pi. 7, fig. 22; pi. 13, fig. 12. 

Range. Two probable examples of this species were found in bed 
474.3, Oxynotum Subzone. 

Genus RADSTOCKICERAS Buckman. 1918 

Remarks. As well as the two species described below, two very 
large fragments of Radstockiceras sp. indet. were found in beds 
544.4 and 544.6; both are about 350 mm diameter and one of them is 
septate up to about 300 mm diameter; a poorly preserved specimen 
was also found in bed 548. 

In their revision of the holotype of Radstockiceras buvignieri, 
Donovan & Guerin-Franiatte (in Fischer, 1994: 68) said that 
Radstockiceras was a late oxynoticeratid that appeared in the 
Jamesoni Zone (from the evidence of Tutcher & Trueman, 1925: 
598, 642) and was not present in the Raricostatum Zone (which was 
the supposed horizon at Radstock of Buckman's (1918: 288) holotype 
of the type species of Radstockiceras). However, the four large 
examples from bed 494 described below are a genuine record of 
Radstockiceras from the Macdonnelli Subzone, Raricostatum Zone, 
whatever may be held to be their specific determination. 



PLATE 5 

Fig. 1 Radstockiceras buvignieri (d'Orbigny). Bed 494, CA 3744. x 0.6; wholly septate. 

Fig. 2 Xipheroceras dudressieri (d'Orbigny). Bed 446.33, C.49336; the asterisk marks the probable end of the phragmocone. 

Fig. 3 Promicroceras planicosta (J. Sowerby ). Bed 45 1 , CA 3927. 

Fig. 4 Radstockiceras sphenonotum (Monke). Bed 544.4, CA 3758; wholly septate. 

Fig. 5 Xipheroceras ziphus (Zieten). Bed 45 1 . CA 3784; wholly septate. 

Fig. 6 Cymbites laevigatus (J. de C. Sowerby). Bed 448.1. CA3763; 6a, b, x 1; 6c, d, x 3; the last septa at the position shown are approximated and 

probably adult. 
Fig. 7 Microderoceras birchi (J. Sowerby). Bed 433.3, CA 3977, x 0.5; a complete (?adult) specimen with a body chamber nearly Wi whorls long. 
Fig. 8 Apoderoceras subtriangulare (Young & Bird). Bed 501.1. CA 3981; a septate fragment. 
Fig. 9 Eoderoceras armatum (J. Sowerby). Neotype. probably from bed 497, C.67323, x 0.75. 
All figures natural size, except Figs 1. 6c, 6d, 7 and 9. 



LOWER LIAS OF ROBIN HOOD'S BAY 



PLATE 5 



131 




132 

Radstockiceras buvignieri (d'Orbigny, 1844) PI. 5, fig. 1 

1844 Ammonites buvignieri d'Orbigny: 261, pi. 74, figs 1-3. 

1855 Ammonites complanosus Simpson: 79, 80. 

1855 Ammonites retentus Simpson: 84. 

1920 Retenticeras retentum (Simpson); Buckman: pi. 166 

(holotype, GSM 26401). 
1962 Metoxynoticeras complanosum (Simpson); Howarth: 105, 

pi. 15, fig. 4 (holotype, WM 239, now lost). 
1992 Radstockiceras complanosum (Simpson); Schlegelmilch: 

60. pi. 54, fig. 2 (WM 239). 
1994 Radstockiceras buvignieri (d'Orbigny); Fischer: 67, pi. 2 1 , 

fig. 3 (holotype, from Breux, Meuse, France). 

Remarks. Occurs in beds 494, Macdonnelli Subzone, 505.2, 
Taylori Subzone, and 544.7, Brevispina Subzone; 7 specimens. 

Measurements (in mm) 





D 


Wh 


Wb 


U 


CA 3744 


167.0 


94.5(0.57) 


42.0 (0.25) 


7.0 (0.04) 


CA 3744 


138.5 


79.0(0.57) 


33.5 (0.24) 


6.3 (0.05) 



REMARKS. The four examples obtained from bed 494 are all large 
specimens of 145-250 mm diameter preserved in grey limestone. 
The best one (PI. 5, fig. 1 ) is wholly septate up to its maximum size 
of 168 mm diameter. The single specimen from bed 505.2 is a well- 
preserved fragment of a part of a whorl, with whorl height 72 mm and 
whorl breadth 27 mm (if the whorl height is 0.57 of the diameter, then 
the whorl breadth is 0.21 of the diameter). One of the two pyritized 
specimens found in bed 544.7 is 25 mm diameter and is similar to the 
holotype of 'Retenticeras' retentum figured by Buckman (1920: pi. 
166). With their very small umbilici (4-5% of the diameter) and 
compressed whorl sections (whorl breadth 21-25% of the diameter), 
all appear to be genuine examples of Radstockiceras buvignieri, of 
which the holotype (Fischer, 1994: 67, pi. 21, fig. 3-178 mm 
diameter, 99 (0.56), 37 (0.2 1 ), 6 (0.03 )) has closely similar characters. 
The holotype of Simpson's species complanosum, and the almost 
identical specimen BM 37960 (from Robin Hood's Bay), undoubt- 
edly belong to the same species, but it is difficult to identify their 
horizons - they could have come from any of the beds 494, 505 and 
544. 



M.K. HOWARTH 

15 specimens; 2 specimens of Cymbites sp. indet. occur slightly 
higher in beds 464.32 and 464.33. 

Remarks. The main species present in Robin Hood's Bay is C. 
laevigatus, but a more compressed species with obsolete ribbing, eg. C. 
fastigatus Schindewolf (1961: 211, pi. 30, figs 8-10), may also be 
present amongst the small specimens determined as Cymbites sp. 
indet. in the Denotatus and Simpsoni Subzones. 



Superfamily EODEROCERATACEAE Spath, 1929 

Family EODEROCERATIDAE Spath, 1929 

Genus MICRODEROCERAS Hyatt, 1871 

Microderoceras scoresbyi (Simpson, 1843) 

1843 Ammonites scoresbyi Simpson: 12. 

1911 Xipheroceras scoresbyi (Simpson): Buckman: pis 39A. B 
(holotype, WM 1 73). C (topotype, GSM 23616), both prob- 
ably from bed 441 .2. 

RANGE. One specimen found in bed 441.2, Birchi Subzone. 

Remarks. This single specimen differs from M. birchi in having 
higher and thicker whorls that are slightly less evolute. 



Microderoceras birchi (J. Sowerby, 1820) 



PI. 5, fig. 7 



1820 Ammonites birchi J. Sowerby: 121, pi. 267. 

1 96 1 Microderoceras birchi (J. Sowerby); Dean et al. : pi. 66, fig. 

3 (BM 67973, from the Dorset coast). 
1973 Microderoceras birchi (J. Sowerby); Donovan & Forsey: 

10. pi. 1, fig. 1 (lectotype, BM 43923, from the Dorset 

coast). 

Range. Found only in bed 433.3, Birchi Subzone; 5 specimens. 

Remarks. The five large specimens in bed 433.3 have typical very 
evolute whorls, with bituberculate ribs up to the end of the largest 
specimen at 235 mm diameter, and considerable variation in whorl 
thickness. The specimen figured in PI. 5. fig. 7 appears to have a 
complete body-chamber nearly 1 Vi whorls long, ending in an ( ?adult ) 
aperture at 210 mm diameter. 



Radstockiceras sphenonotum (Monke, 1888) PI. 5, fig. 4 

1888 Ammonites sphenonotus Monke: 228. pi. 2/3, fig. 14 

(holotype, from Germany). 
1914 Oxynoticeras sphenonotum (Monke); Pia: 65, pi. 7. fig. 12. 

Range. Beds 542.1-544.4, Polymorphus Subzone; 8 specimens. 

Remarks. The best preserved specimen occurs in bed 544.4 (PI. 5, 
fig. 4). 



Family CYMBITIDAE Buckman, 1919 
Genus CYMBITES Neumayr, 1979 



Cymbites laevigatus (J. de C. Sowerby, 1827) PI. 5, fig. 6 

1827 Ammonites laevigatus J. de C. Sowerby: 135, pi. 570, fig. 3. 
1957 Cymbites laevigatus (J. de C. Sowerby); Donovan: 413, figs 

1-8 (topotypes (the holotype is lost), from Brooki to Stellare 

Subzones, Dorset coast). 

Range. Beds 446.5^464.1, top Obtusum to Simpsoni Subzones; 



Genus XIPHEROCERAS Buckman, 1911 

Xipheroceras dudressieri (d'Orbigny, 1845) PI. 5, fig. 2 

1845 Ammonites dudressieri d'Orbigny: 325. pi. 103, figs 1, 2. 
1926b Xipheroceras dudressieri (d'Orbigny); Spath: 172, pi. 9. 

fig. 6 (BM C. 2235a, a typical example from the Obtusum 

Subzone, Dorset). 
1994 Xipheroceras dudressieri (d'Orbigny); Fischer: 91, pi. 19, 

fig. 3 (from Mulhausen. France). 



Range. One specimen in bed 446.33, Obtusum Subzone. 



Xipheroceras ziphus (Zieten, 1830) 
1830 



PI. 5 fig. 5 



Ammonites ziphus Zieten: 6. pi. 5, fig. 2 (holotype, BM 

62590. from Heiningen, Wurttemberg, Germany). 
1926 Xipheroceras ziphus (Zieten); Buckman: pi. 732 (GSM 

47832, coarsely ribbed inner whorls, from the Obtusum 

Subzone, Dorset). 
1928 Xipheroceras revertens Buckman: pis 772 A, B (a typical 

large example of X. ziphus from the Obtusum Subzone. 

Dorset). 



LOWER LIAS OF ROBIN HOOD'S BAY 



133 



1973 Xipheroceras planicosta Buckman; Donovan & Forsey: 9, 
pi. 3, figs 1, 2 (GSM 25033, Obtusum Zone, Dorset). 

Range. Beds 446.4^153.1, Obtusum and Stellare Subzones; 7 
specimens. 

Remarks. In addition to those listed above, single examples of 
Xipheroceras sp. indet. occur in beds 446.32, 453.2 and 453.3. 

Genus BIFERICERAS Buckman, 1913 

Bifericeras bifer (Quenstedt, 1845) 

1845 Ammonites bifer Quenstedt: Quenstedt: 83, pi. 4, fig. 14 

(from Wiirttemberg, Germany). 
? 1925a Ophidewceras ziphoides Spath: 138^0, figs 1, 2a, 2b 

(from low in the Oxynotum Subzone, Mill Beck Nab, Robin 

Hood's Bay; originally in Hull Museum, but now destroyed; 

see Donovan & Forsey, 1973: 16). 
1957 Bifericeras bifer (Quenstedt); Soil: 402, pi. 19, figs 1-7 

(from Wiirttemberg, Germany). 
1976 Bifericeras bifer (Quenstedt); Schlegelmilch: 58, pi. 25, fig. 

3 ('neotype', from Wiirttemberg, Germany). 
1990 Bifericeras bifer (Quenstedt); Hollingworth et al: 165, pi. 

2, figs 1-12 (BM C.93398-93409, from the Oxynotum 

Subzone, Somerset). 

Range. Bed 483.1, Oxynotum Subzone; 5 specimens. The de- 
stroyed holotype of Ophidewceras ziphoides Spath might have been 
a large example of Bifericeras bifer. 

Bifericeras vitreum (Simpson, 1855) 

1855 Ammonites vitreus Simpson: 46. 

1924 Microceras vitreum (Simpson); Buckman: pi. 529 (holotype, 

WM 462, possibly from bed 486.2). 
1976 Bifericeras vitreum (Simpson); Schlegelmilch: 146, pi. 25, 

fig. 9 (WM 462). 

RANGE. One specimen in bed 486.2, Oxynotum Subzone. 

Bifericeras donovani Dommergues & Meister, 1992 

PL 8, fig. 3 

1992 Bifericeras donovani Dommergues & Meister: 223, figs 
5(8)-5(10), figs 7(1), 7(2), 7(3) (holotype), 7(4)-7(l 1) (all 
from nodules in the lower part of bed 501 . 1 at Wine Haven). 

Range. Occurs only in bed 501.1, base of Taylori Subzone; 18 
specimens. 

Genus CRUCILOBICERAS Buckman, 1920 



Crucilobiceras densinodulum Buckman, 1923 



1876 



PI. 6, fig. 1 



Aegoceras obsoletum (Simpson); Blake: 276, pi. 7, fig. 1 

(BMC. 17939, from bed 486.3). 
1923 Crucilobiceras densinodulum Buckman: pi. 442 (holotype, 

from Lyme Regis, Dorset). 
1926/? Crucilobiceras ornatilobatum Spath: 176, pi. 11, fig. 1 

(holotype, from Lyme Regis, Dorset). 

Range. Beds 486.3-488, Densinodulum and Raricostatoides 
Subzones; 19 specimens. 

Remarks. The well-preserved C. densinodulum in beds 486.3 (PI. 
6, fig. 1 ) are unituberculate (ie. have ventro-lateral but no prominent 



umbilical-lateral tubercles), are moderately to finely ribbed on the 
inner whorls, and are characteristically much more compressed than 
C. densinodum (Oppel). Small specimens or inner whorls of C. 
densinodulum are very similar to 'Bifericeras (Hemimicroceras)' 
subplanicosta (Oppel, 1856) and the relationship between the two 
(eg. a macroconch/microconch pair or size difference only) has yet 
to be resolved. 

Genus EODEROCERAS Spath, 1925 

Eoderoceras armatum (J. Sowerby, 1815) 

PI. 5, fig. 9; PI. 8, fig. 2 

1815 Ammonites armatus J. Sowerby: 215, pi. 95. 

1843 Ammonites anguiformis Simpson: 17. 

1843 Ammonites owenensis Simpson: 25. 

1855 Ammonites miles Simpson: 65. 

1880/82 Aegoceras armatum (J. Sowerby); Wright: 340 (1882). pi. 
28 (1880), figs 1, 2 (SM J18222), 3-5 (SM J18223), both 
from beds 497-99; non pi. 29, =Eoderoceras pugnax (Buck- 
man, 1914: 103c). 

1911 Deroceras miles (Simpson); Buckman: pi. 44 (holotype, 
WM 162, from bed 498 or 499). 

1912 Deroceras anguiforme (Simpson); Buckman: pi. 64 
(holotype, WM 86, from beds 497-99). 

1912 Deroceras owenense (Simpson); Buckman: pi. 65 (holotype, 

WM 476, from beds 497-99). 
1926£> Deroceras eusculptum Spath: 175, pi. 10, fig. 3 (BM 

C. 26907, from Lyme Regis, Dorset). 
1992 Eoderoceras gr. miles (Simpson); Dommergues & Meister: 

231, figs 5(5), 5(7) (from bed 497). 
2000 Eoderoceras armatum (J. Sowerby); Blau et al: 269, figs 4 

( 1-7), 5 ( 1, 3, 5), 6 ( 1), 7, 8 (from NW Germany). 

Range. Beds 494-499, Macdonnelli and Aplanatum Subzones; 
62 specimens. 

Type specimen. BM C.67323, Sowerby Collection, from Robin 
Hood's Bay, probably from bed 497, is here designated neotype. 

Measurements (in mm) 





D Wh 


Wb 


U 


BM C.67323 


128.0 32.0(0.25) 


31.5(0.25) 


69.8 (0.54) 


BM C.67323 


102.0 27.4(0.27) 


- 


53.3 (0.52) 



Remarks. Sowerby (1815: 2 15) stated that many examples of his 
species, including the specimen that he figured, which was collected 
by Mr. Strangewayes, came from 'the great Alum-clay formation at 
Whitby' (a stratigraphical term that was used by Sowerby, Young & 
Bird, Sedgwick and others for beds that included ones as low as the 
Upper Sinemurian/Lower Pliensbachian clays). Sowerby did not 
mention any other locality in his original description, so it is not 
possible to agree with Spam's (1925a: 137, 167) statements that 'the 
true D. armatum apparently does not occur in Yorkshire', and 
'Sowerby's type is not preserved in the BM . . . and has always 
seemed to me to be more like a Charmouth than a Whitby specimen' , 
nor with Donovan's (1958: 32) opinion that 'Sowerby's type speci- 
men . . . was almost certainly obtained from the Dorset coast'. It is 
not permissible to transfer the type locality to Dorset on the opinion 
that the Yorkshire specimens are morphologically different from 
those in Dorset, or the belief that the pyritic preservation and 
frequency of decay perceived from Sowerby's figure are more 
reminiscent of Dorset examples. The Yorkshire specimens are not 
morphologically different from those in Dorset (two especially fine 



134 



PLATE 6 



M.K. HOWARTH 




LOWER LIAS OF ROBIN HOOD'S BAY 



135 



Yorkshire specimens had been figured by Wright, 1 880: pi. 28), and 
there are many Yorkshire examples preserved in iron pyrites, espe- 
cially in beds 497 and 499, which are also subject to decay, though 
less readily than the Dorset ones. 

After Spath wrote about the species in 1925, the remaining 
ammonites in the Sowerbys' collections were obtained by The 
Natural History Museum in 1935, and amongst them is a medium to 
large example (BM C. 67323) of Eoderoceras armatum from Robin 
Hood's Bay, which might even have been one of Sowerby's original 
syntypes. At 132 mm diameter it is slightly larger than Sowerby's 
figure of a 117 mm diameter specimen (assuming that his figure is 
natural size), but it does not differ in any morphological feature from 
that figure, and the part now missing at the beginning of the final 
whorl was once present, judging from remaining traces of old glue. 
There are some patches of pyritic preservation on the inner whorls, 
which fortunately do not appear to be subject to decay. As a possible 
original syntype, and a definite topotype, it is a close morphological 
match with Sowerby's figured specimen and is clearly the best 
neotype that can now be selected. It presents an unexpected opportun- 
ity to finally settle the identity of this frequently quoted species. 

The neotype (PI. 5, fig. 9) consists of 6-6'/2 septate whorls up to 
about 110 mm diameter, followed by a quarter of a whorl of body 
chamber ending at 1 32 mm diameter. The whorls are very evolute, the 
whorl section is near-circular and the umbilical wall and edge are 
evenly rounded. There are many fine indistinct radial ribs between 
stronger periodic lateral ribs that end in prominent ventro-lateral 
tubercles. Ribs of moderate strength cross the venter, curving gently 
forwards, and there are 3-5 such ribs between adjacent ventro-lateral 
tubercles. There are no umbilical tubercles. 

In Robin Hood's Bay there are a few specimens in the Macdonnelli 
Subzone, then the species becomes much more common in the 
Aplanatum Subzone, with 44, 8 and 7 examples collected by Bairstow 
from beds 497, 498 and 499 respectively. The highest in bed 499 is 0. 1 5 
m below the top, and only 0.23 m below the top of the Sinemurian. Six 
specimens were found lower in bed 499 (0. 15-0.37 m above the base), 
and the best preserved of them is figured in PI. 8, fig. 2; the latter has a 
phragmocone ending at ca. 68 mm diameter at the position indicated on 
the figure, then it has a body-chamber 1.125 whorls long ending at a 
final aperture at ca. 125 mm diameter (the final 0.35 whorls are 
detached and poorly preserved, and are not figured). There is no 
overlap between Eoderoceras and Apoderoceras, and the lowest 
example of the latter genus occurs in bed 501 . 1 just 0.36 m above the 
highest Eoderoceras. 

Eoderoceras hastatum (Young & Bird, 1828) PI. 6, fig. 3 

1828 Ammonites hastatus Young & Bird: 261, pi. 14, fig. 3. 
1914 Deroceras hastatum (Young & Bird); Buckman: pis 102A, 

B (?holotype, WM 661, from bed 493.2). 
1914 Deroceras impavidum Buckman: pi. 104 (holotype, WM 

166; probably from bed 493.2). 

Range. Found only in bed 493.2, Raricostatoides Subzone; 4 
specimens. 



Remarks. E. hastatum has more depressed whorls and more 
widely spaced ventro-lateral tubercles than E. armatum which occurs 
at higher levels. The specimen figured here (PI. 6, fig. 3) is very like 
the holotype in having striate ribs angled strongly backwards from 
the umbilical edge, and the almost identical holotype of Buckman's 
species impavidum probably came from the same bed 493.2. 

Other species of Eoderoceras: 

lE.diversum (Simpson, 1843: 13); Blake, 1876: 282, pi. 8, fig. 3 (SM 
J34799); Howarth, 1962: 107, pi. 15, fig. 9 (SMJ34799, neotype). 
The neotype represents a highly evolute, serpenticone species, 
which might be an Eoderoceras, but no specimens were found by 
Bairstow. 

Genus PROMICROCERAS Spath, 1925 

Promicroceras planicosta (J. Sowerby, 1814) PI. 5, fig. 3 

1814 Ammonites planicosta J. Sowerby: 167, pi. 73. 

1822 Ammonites aureusYoung & Bird: 248, pi. 13, fig. 6) (type 

specimen lost). 
71843 Ammonites siphuncularis Simpson: 46. 
71912 Androgynoceras siphunculare (Simpson); Buckman: pi. 48 

(holotype, WM 485, from beds 451-454). 
1925a Promicroceras planicosta (J. Sowerby); Spath: 299-302, 

fig. 8f. 
1925a Promicroceras aureum (Young & Bird); Spath: 301 , fig. 8d 

(BM 17160, possibly from 451). 
1926ft Promicroceras planicosta (J. Sowerby); Spath: 171, pi. 9, 

figs 1 (BM C.26337), 7 ('neotype', BM C.2235b); both 

from Charmouth, Dorset. 

Range. Beds 446.33-454. 1 , Obtusum and Stellare Subzones; 290 
specimens. 

Remarks. The current interpretation of Promicroceras planicosta 
may not be satisfactory. After lengthy discussion, Spath (1925a: 
299-302) selected as neotype the specimen BM C.2235b (T. Wright 
Colin, 1887) from Charmouth, Dorset (almost certainly from bed 
85), even though Sowerby (1814: 167) said that his main specimens 
came from Marston Magna, and it is highly probable that the original 
block of specimens that he figured (Sowerby, 1814: pi. 73, now lost) 
came from the Marston Marble at Marston Magna, Somerset. Spath 
(1925a: 305) then created a new species, P. marstonense, for the 
form at Marston Magna, using as holotype a specimen (BM 43914b) 
from Sowerby's syntypes of P. planicosta. The selection of a 
Charmouth specimen as neotype of P. planicosta was unfortunate, 
and may be invalid because there were Marston Magna specimens 
available amongst the original syntypes. The designation of a Marston 
Magna specimen as neotype (or lectotype) would have been much 
more in accordance with Sowerby's original concept of his species. 
In any case, the forms of Promicroceras at Charmouth and Marston 
Magna appear to be very close and the two names are probably 



PLATE 6 

Fig. 1 Crucilobiceras densinodulum Buckman. Bed 486.3. CA 3828; the body chamber is exactly one whorl long. 

Fig. 2 Apoderoceras aculeatum (Simpson). Bed 526.5, CA 4031. 

Fig. 3 Eoderoceras hastatum (Young & Bird). Bed 493.2, CA 3895, x 0.6; suture-lines that probably mark the end of the phragmocone are visible exactly 

one whorl before the aperture. 
Figs 4, 5 Apoderoceras subtriangulare (Young & Bird). 4, bed 520.7, CA 4018; probably a complete adult microconch, with a body chamber exactly one 

whorl long and a slightly contracted final aperture. 5, bed 502, CA3990; the asterisk marks the probable end of the phragmocone. 
All figures natural size, except Fig. 3. 



136 



M.K. HOWARTH 



synonyms. Two other names that are probably also synonyms of P. 
planicosta are P. perplanicosta (Spath, 1925a: 269; 1926£>, 172, pi. 
9, fig. 2) and P. precompressum Spath ( 1926b, 173, pi. 9, fig. 5), both 
from beds 83 or 85 at Charmouth. 

Promicroceras capricornoides (Quenstedt, 1883) 

1883 Ammonites capricornoides Quenstedt: 129, pi. 17, fig. 11 

(from Wtirttemberg, Germany). 
1926i> Promicroceras capricornoides (Quenstedt); Spath: 172, pi. 

9, fig. 3 (from the Turneri Zone, Charmouth, Dorset). 

Range. Beds 436^446.32, Birchi and basal Obtusum Subzones; 
39 specimens. 

REMARKS. P. capricornoides is slightly more involute, has a more 
rapidly increasing whorl height and slightly fewer ribs than P. 
planicosta. 



Family COELOCERATIDAE Haug, 1910 
Genus APODEROCERAS Buckman, 1921 



REMARKS. Identification of the horizons from which the Yorkshire 
type and figured specimens were obtained is especially uncertain in 
Apoderoceras. The originals of A. subtriangulare and its two syno- 
nyms (A. hamiltoni and A. spicatum) could have come from any of 
beds 502, 504-07 509, 520 and 522, all of which contain many large 
fragments of outer whorls as well as a few more complete specimens 
like the original of A. 'hamiltoni'. Similarly, the originals of A. 
aculeatum and its three synonyms (A. decussation, A. mutatum and 
A. leckenbyi) could have come from any of beds 523-526. 

Apoderoceras subtriangulare (Young & Bird, 1822) 

PI. 5, fig. 8; PI. 6, figs 4, 5 

1 822 Ammonites subtriangularisYoung & Bird: 250. pi. 12, fig. 4. 
1843 Ammonites hamiltoni Simpson: 27. 
1843 Ammonites spicatus Simpson: 28. 

1913 Deroceras subtriangulare (Young & Bird); Buckman: pi. 
71 (holotype, WM927). 

1914 Deroceras spicatum (Simpson); Buckman: pi. 103 
(holotype, WM 920). 

1924 Apoderoceras hamiltoni (Simpson); Buckman: pls530A,B 

(both are the holotype, WM 165). 
1992 Apoderoceras sp. indet.; Dommergues & Meister: 232, fig. 

5(6) (from nodules at the middle of bed 50 1 . 1 , Wine Haven). 

Range. Beds 501.1-522.1, Taylori Subzone; 41 specimens. 

Remarks. The lowest examples of Apoderoceras are three speci- 
mens in bed 501.1. Two are very small cadicones, one large enough 
to have ventro-lateral tubercles, while the third (PI. 5, fig. 8) is part of 
one side of a whorl at a whorl height of 35 mm that has obscure radial 
ribs and striae and pointed ventro-lateral tubercles. It is identical 
with A. subtriangulare at a similar size, and is important for fixing 
the base of the Taylori Subzone at this level. 

Large A. subtriangulare up to 350 mm diameter, with large 
ventro-lateral spines and broad flat venters occur in bed 502 and at 
many more horizons up to bed 522.1. Most are fragments of outer 
whorls with nothing to link them to the few inner whorls, but an 
example in bed 507.1 (CA 4006) has both inner and outer whorls, 
and proves that small inner whorls like those in PL 6, fig. 5 and the 
very well-preserved specimen of PL 6, fig. 4 develop into the massive 
outer whorls with broad flat venters and very large ventro-lateral 



spines that are characteristic of A. subtriangulare. The holotype of A. 
subtriangulare (Buckman, 1913: pi. 7 1) is a fragment of such a large 
outer whorl. Of the two Simpson species that are placed in syn- 
onymy, the holotype of A. spicatum (Buckman, 1914: pi. 103) is a 
very similar fragment of a large outer whorl, though the broad flat 
venter is not shown because only one side is preserved, while the 
holotype of A. hamiltoni (Buckman, 1924: pi. 530) is one of the few 
specimens that show inner and outer whorls preserved in the same 
individual. [Note the remarkable resemblance of PL 6, fig. 4 to a 
depressed-whorled species of the Toarcian genus Peronoceras. eg. P. 
perarmatum (Young & Bird), Howarth, 1978: pi. 4, fig. 7; the latter 
differs only in having well-defined ribs on the side of the whorl, 
compared with the poor, irregular or striate ribs on the inner whorls 
of Apoderoceras subtriangulare]. 

Apoderoceras aculeatum (Simpson, 1843) 

PI. 6, fig. 2: PI. 7, fig. 1 

71843 Ammonites marshallani Simpson: 24. 

1843 Ammonites decussatus Simpson: 25. 

1843 Ammonites aculeatus Simpson: 27. 

1855 Ammonites mutatus Simpson: 63. 

1876 Aegoceras aculeatum (Simpson); Blake: 278, pi. 7, fig. 4 
(BMC. 17878). 
1880/82 Aegoceras leckenbyi Wright: 344 (1882), pi. 30 (1880), 
figs 1-3 (lectotypeof/ecfcenbv/ (designated Howarth, 1962: 
109), SM J 18224), figs 4-7 (paralectotype of leckenbyi, 
also holotype of decussation. SM J 18225). 

1913 Deroceras aculeatum (Simpson); Buckman: pis 72A-C 
(paratype, WM 177; the holotype is lost). 

1914 Deroceras mutatum (Simpson); Buckman: pi. 105 (holotype, 
GSM 26406). 

1962 Apoderoceras decussatum (Simpson); Howarth: 109, pi. 
16, fig 1 (holotype, SM J18225). 
71962 Apoderoceras marshallani (Simpson); Howarth: 109, pi. 
15, fig. 5 (holotype. WM 468); the holotype is only 16 mm 
diameter, and although it could be a very small example of 
A. aculeatum. Simpson's species is not accurately determin- 
able. 

Range. Beds 523-526.6, Taylori Subzone; 1 5 specimens. There are 
also single specimens of Apoderoceras sp. indet. in beds 526.7 and 529. 

Remarks. Specimens from beds 523-526 differ from A. 
subtriangulare in beds 501-522 in having the outer whorls more 
rounded, with evenly arched whorl sides and venter, no definite ventro- 
lateral angle, and smaller ventro-lateral tubercles on the inner whorls. 
These stratigraphically higher specimens belong to the species A. 
aculeatum (Simpson), of A. decussatum and A. leckenbyi are synonyms 
(Ammonites aculeatum and A. decussatum are both of Simpson, 1843, 
and therefore have equal priority; the name for the species was selected 
by Blake (1876: 278), who, as 'First Reviser' (ICZN Code, art. 24.2), 
chose aculeatum as the name for the species). A. marshallani. another 
Simpson name of the same date ( 1 843), is probably also a synonym, but 
the holotype is too small to be definitely identifiable, and A. mutatum 
(Simpson. 1855)andA. leckenbyi (Wright, 1880) are two more syno- 
nyms, judging from their well-preserved type specimens. 

The specimen figured in PL 7, fig. 1 is a fairly large, wholly septate 
example of A. aculeatum and PL 6, fig. 2 has well-preserved inner 
whorls showing much smaller ventro-lateral tubercles than in A. 
subtriangulare (cf. inner whorls of PL 6, fig. 5). 

Other species of Apoderoceras: 

1. lApoderoceras sinuatum (Simpson, 1855: 62); Buckman, 1914: 



LOWER LIAS OF ROBIN HOOD'S BAY 



137 



pi. 94 (holotype, WM 160). No specimens resembling this 
holotype were found by Bairstow. 
2. 1A. armiger (Simpson, 1855: 66 (non Ammonites armiger J. deC. 
Sowerby, 1840)); Howarth, 1962: 107. From Simpson's descrip- 
tion this was probably an Apoderoceras, but the type specimen is 
lost and the species is not identifiable. 

Genus HYPERDEROCERAS Spath, 1926 

Remarks. Bairstow found only one poorly preserved example (CA 
4053) of Hyperderoceras sp. indet. in bed 540.1 (Polymorphus 
Subzone). This shows parts of inner whorls at 1 5-50 mm diameter that 
have strong ribs and ventro-lateral tubercles, followed by septate 
fragments of larger whorls with a quadrate whorl section, where both 
whorl height and breadth are 50-60 mm. ie. they are not compressed 
like Epideroceras at this size. Any of the following four Simpson 
species might have come from this horizon, but the Bairstow specimen 
is not specifically identifiable: 

1. Hyperderoceras niamillatum (Simpson. 1843: 28 (non Ammo- 
nites mamillatus Schlotheim, 1813)); Howarth, 1962: 108, pi. 15, 
fig. 6 (neotype (designated by Howarth, 1962: 108), WM 2102). 

2. H. validum (Simpson, 1855: 39); Blake, 1876: 278, pi. 7, fig. 3 
(SM-?holotype): Buckman, 1913: pi. 83 (holotype, SM J3275). 

3. H.retusum (Simpson, 1855: 62); Buckman, 1913: pi. 82 (holotype, 
WM 184). 

4. H.nativum (Simpson. 1855: 68); Buckman, 1913: pi. 84 (holotype, 
WM931). 



Family PHRICODOCERATIDAE Spath, 1938 
Genus PHR1CODOCERAS Hyatt, 1900 

Remarks. Species of Phricodoceras show a considerable amount 
of variation in rib-density, in the development of lateral tubercles, 
and in overall size of the tubercles. The specimens from Robin 
Hood's Bay suggest that a sparsely-ribbed species, here identified as 
P. taylori, occurs in the lower half of the Taylori Subzone, while a 
more densely-ribbed species, P. cornutum, occurs in the upper half of 
the subzone. Tubercle strength depends to some extent on preserva- 
tion, and in any species tubercles are more prominent when the shell 
is preserved, compared with their appearance on internal moulds. 
The Robin Hood's Bay specimens are identified below mainly 
according to their rib-density, but tubercles are accorded some 
significance when they are especially large. 

Phricodoceras taylori (J. de C. Sowerby, 1826) 

1826 Ammonites taylori J. de C. Sowerby: 23, pi. 514, fig. 1 
(holotype, lost (originally in Norwich Museum), from Gla- 
cial Drift. Happisburgh, north Norfolk, perhaps derived 
from the Yorkshire coast). 

1855 Ammonites quadricornutus Simpson: 71. 

1884 Ammonites quadricornutus Simpson: 106. 

1911 Phricodoceras quadricornutum (Simpson); Buckman: pi. 
33 (holotype, WM 495 ; possibly from beds 501-517, lower 
than P. cornutum). 

RANGE: Beds 50 1 .3-524. 1 , Taylori Subzone; 8 specimens. 

Remarks. P. taylori occurs in the lower and middle parts of the 
Taylori Subzone at Robin Hood's Bay, where the poorly preserved 
specimens have the widely spaced ribs (12-13 per whorl at 50 mm 
diameter) that are typical of the species. The holotype of Ammonites 
| quadricornutus has the same rib-density, and the strength of the 



tubercles is similar to that shown in Sowerby's figure of taylori, even 
after allowances are made for slightly different modes of preservation. 

Phricodoceras cornutum (Simpson, 1843) PI. 7, fig. 2 

1843 Ammonites cornutus Simpson: 31. 

1 855 Ammonites cornutus Simpson: 7 1 . 

1 884 Ammonites taylori J de C. Sowerby; Simpson: 105 (includ- 
ing Ammonites cornutus which Simpson now considered to 
be a synonym). 

1911 Phricodoceras cornutum (Simpson); Buckman: pi. 32 
(holotype, WM 185). 

1976 Phricodoceras cornutum (Simpson); Schlegelmilch: 152, 
pi. 28, fig. 1 (WM 185). 

RANGE: Beds 524.3-530.2, Taylori Subzone; 8 specimens. 

Remarks. P. cornutum occurs higher in the Taylori Subzone than P. 
taylori, and differs from the latter in having more ribs ( 1 7-20 per whorl 
at 50 mm diameter) and smaller tubercles. 

Phricodoceras nodosum (Quenstedt, 1846) 

1846 Ammonites taylori nodosus Quenstedt: 136, pi. 9, fig. 21 

(holotype, from Wurttemberg, Germany) (non Ammonites 

nodosus, Bruguiere, 1789). 
1961 Phricodoceras aff. taylori (J. de C. Sowerby); Dean et al.\ 

pi. 68, fig. 5 (BM C. 1 798 1 . probably from bed 520). 
1980 Phricodoceras nodosum (Quenstedt); Schlatter: 78. pi. 6, 

figs 5. 6 (from Wurttemberg. Germany). 

Range. A single specimen in bed 520.5, Taylori Subzone. 

REMARKS. All the specimens listed above have the same rib-density 
as P. taylori, but they have much more prominent tubercles. Even on the 
internal mould the tubercles appear to be genuinely much larger, as can 
be seen on the specimen figured by Dean et al (\96l), which has both 
shell and internal mould preserved on different portions of the shell. 

Genus EPIDEROCERAS Spath. 1923 

Remarks. The only examples of Epideroceras found by Bairstow 
were parts of two specimens in bed 542.4, Polymorphus Subzone. 
They are fragments of large septate whorls, with compressed whorl 
sections at 45-60 mm whorl height, a rounded venter, and radial ribs, 
but no tubercles. They are too fragmentary and poorly preserved to be 
specifically identified. The following Simpson species might have 
come from bed 542.4: 

Epideroceras sociale (Simpson, 1855: 39); Blake, 1876: 278, pi. 7. 
fig. 6 (SM collection); Buckman, 1914: pi. 95 (holotype, WM 68). 



Family POLYMORPHITIDAE Haug, 1887 
Genus GEMMELLAROCERAS Hyatt, 1900 

Synonyms. Tubellites Buckman, 1924; Leptonotoceras Spath, 
1925. 

REMARKS. Gemmellaroceras has been divided in two subgenera: 
Gemmellaroceras s.s., in the Jamesoni Zone and younger beds, in 
which the first lateral lobe is trifid, and an earlier Raricostatum to basal 
Jamesoni Zone subgenus G. (Leptonotoceras), in which the first lateral 
lobes are bifid (Dubar & Mouterde, 1961: 237; Geczy, 1976: 73-75). 
In Robin Hood's Bay Gemmellaroceras ranges from the top part of the 
Macdonnelli Subzone up to near the top of the Taylori Subzone. Most 
specimens belong to the very small species G tubellum (Simpson) (the 
type species of Tubellites Buckman, 1924), and those specimens that 



PLATE 7 



M.K. HOWARTH 




LOWER LIAS OF ROBIN HOOD'S BAY 



139 



are large enough to have divided first lateral lobes appear to have the 
bifid lobes of Leptonotoceras. However, many of the specimens are 
very small and the largest (in bed 501 .2) is only 19 mm diameter. In the 
top part of the Taylori Subzone at Robin Hood's Bay the much larger 
species Gemmellawceras rutilans has the trifid first lateral lobes of 
Gemmellaroceras s.s. Whether this division into subgenera is real, oris 
a function of the size to which species grow and therefore the complex- 
ity attained by their suture-lines, has still to be resolved. 

Gemmellaroceras tubellum (Simpson, 1855) 

1855 Ammonites tubellus Simpson: 42. 
1876 Aegoceras tubellum (Simpson); Blake: 279. pi. 5, fig. 7. 
1924 Tubellites tubellus (Simpson); Buckman: pi. 491 (holotype, 
WM 98 1 ; ?probably from bed 497 or 498). 

RANGE. Beds 495.7-505.1, Macdonnelli to Taylori Subzones; 104 
specimens. 

Range. After its first appearance in the top bed of the Macdonnelli 
Subzone, G. tubellum becomes much more common in the Aplanatum 
and bottom of the Taylori Subzones. 

Gemmellaroceras rutilans (Simpson, 1843) PI. 7, fig. 4 

1843 Ammonites rutilans Simpson: 45. 

1962 Polymorphites rutilans (Simpson); Howarth: 110.pl. 15, 
figs 7 (neotype (designated Howarth, 1962: 1 10), WM 95), 
8 ( WM 94); both probably from bed 526. 1 or 530. 1 . 

RANGE. Beds 526. 1 and 530. 1 . Taylori Subzone; 4 specimens. 

Remarks. G. peregrinum (Haug, 1887: 1 14, pi. 4, fig. 5) is a very 
similar (?or identical) species that occurs in the upper part of the Taylori 
Subzone in Dorset (bed 108), where specimens attain large sizes of up 
to 90 mm diameter. 

Genus POLYMORPHITES Haug, 1887 



Polymorphites caprarius (Quenstedt, 1856) 
1856 



PI. 7, fig. 9 



Ammonites caprarius Quenstedt: 131, pi. 16, fig. 1 (holotype, 

from Balingen, Wurttemberg, Germany; BM C. 55358 is a 

cast of this specimen). 
1885 Ammonites caprarius Quenstedt: Quenstedt: 244, pi. 30, 

figs 40, 41 (holotype refigured). 
1976 Platypleuroceras caprarium (Quenstedt); Schlegelmilch: 

63, pi. 29, fig. 5 (holotype refigured). 
1980 Polymorphites caprarius (Quenstedt); Schlatter: 92. 

Range. Beds 538-542.5, Polymorphus Subzone; 87 specimens. 



Remarks. There seems to be no doubt that Quenstedt's figure of 
1885 (pi. 30. fig. 41) is a drawing ofthe same specimen that he figured 
in 1856 (pi. 16, fig. 1). This specimen (Geological Institute, Tubingen 
University. Ce 5/30/41 ) was figured again by Schlegelmilch (1976); it 
is the holotype. Schlegelmilch's (1976: 63) designation of this speci- 
men as neotype was not necessary. One of Bairstow's better preserved 
specimens from bed 539 is figured in PI. 7, fig. 9. 



Polymorphites trivialis (Simpson, 1843) 

1843 
1876 

1912 



PI. 7, fig. 7 

Ammonites trivialis Simpson: 10. 

Amaltheus trivialis (Simpson); Blake: 292, pi. 5, figs 6a 

(BM C. 17891), 6b-d. 

Polymorphites trivialis (Simpson); Buckman: pi. 53, figs 1, 

la, lb (the lectotype, WM 105, now lost), figs 2, 3 

(paralectotypes). 

Range. Beds 542.4—546.3, Polymorphus and Brevispina Subzones; 
84 specimens. 

REMARKS. Many examples of P. trivialis are found in beds 544.35- 
544.9 at the top of the Polymorphus and bottom of the Brevispina 
Subzones, and it is highly probable that the type specimens came from 
544.35, 544.4, 544.5 or 544.9. 



Polymorphites bronni (Roemer, 1836) 



PI. 7, fig. 5 



1836 Ammonites bronni Roemer: 181, pi. 12, fig. 8 (holotype, 

from north Germany). 
1884 Ammonites bronni Roemer; Quenstedt: 245, pi. 30, figs 44, 

46, 48 (from Wurttemberg, Germany). 
1976 Polymorphites bronni (Quenstedt); Schlegelmilch: 62, pi. 

28, fig. 8 (original of Quenstedt, 1885: pi. 30, fig. 48). 
1980 Polymorphites bronni (Quenstedt); Schlatter: 82, pi. 7, fig. 

1, pi. 11, fig. 5 (from Wurttemberg, Germany). 

RANGE. Beds 554—560.3, Jamesoni Subzone; 36 specimens. 

REMARKS. Occurs in the upper half of the Jamesoni Subzone, and it 
differs from P. trivialis and P. polymorphus in having consistently 
stronger ribbing, small ventro-lateral tubercles and a mid- ventral keel. 
The Robin Hood's Bay specimens have been identified from Schlatter's 
(1980: 82) interpretation ofthe species. 

Polymorphites polymorphus (Quenstedt, 1845) PI. 7, fig. 3 

1845 Ammonites polymorphus quadratus Quenstedt: 87, pi. 4, 
fig. 9 (lectotype. from Germany, designated by Donovan & 
Forsey, 1973: 12). 

1961 Polymorphites polymorphus (Quenstedt); Dean el al.\ pi. 
68, fig. 4 (from Gloucestershire). 



!10 mm diameter. 



PLATE 7 

Fig. 1 Apoderoceras aculeatum (Simpson). Bed 524.3, CA4022. x 0.5; wholly septate up to the aperture at ca. 

Fig. 2 Phricodoceras comutum (Simpson). Bed 525. CA 4070. 

Fig. 3 Polymorphites polymorphus (Quenstedt). Bed 555, CA 43 1 7. 

Fig. 4 Gemmellaroceras rutilans (Simpson). Bed 526.1, CA 4178; approximated adult septa occur at the position marked three-quarters of a whorl before 

the aperture. 
Fig. 5 Polymorphites bronni (Roemer). Bed 560.3 (below the top 0.08 m). CA 4226: wholly septate. 

Platypleuroceras aureitm (Simpson). Bed 546.2, CA 4480; the body chamber appears to be exactly one whorl long. 
Polymorphites trivialis (Simpson). Bed 544.35, CA 4326; wholly septate. 
Aegoceras (A.) maculatum (Young & Bird) var. atavum Spath. Bed 583.2, C. 38874. 

Polymorphites caprarius (Quenstedt). Bed 539, CA 4237; a complete microconch, with a body chamber about two-thirds of a whorl long. 
Tropidocerasfuttereri Spath. 10, bed 560.3 (in the top 0.08 m), CA 4544. 11, bed 568, CA 4545. 
Aegoceras (A.) maculatum (Young & Bird). 12, bed 581. C.41307. 13, bed 590.61, C. 38883. 



Fig. 6 

Fig. 7 

Fig. 8 

Fig. 9 

Figs 10, 11 

Figs 12, 13 

Fig. 14 Aegoceras (Oistoceras) sinuosiforme Spath. Bed 598. 1 , C. 38930. 

All figures natural size, except Fig. 1. 



140 



M.K. HOWARTH 



1973 Polymorphites polymorphus (Quenstedt); Donovan & 

Forsey: 11, 12. 
1980 Polymorphites polymorphus (Quenstedt); Schlatter: 84, pi. 

7, fig. 2 (from Wiirttemberg, Germany). 

RANGE. Found only in bed 555, Jamesoni Subzone; 2 specimens. 

Remarks. Although the lectotype of P. polymorphus, as validly 
designated by Donovan & Forsey (1973: 12), is probably lost, 
Schlegelmilch's (1976: 61) designation of a 'neotype' is not valid 
because it is radically different in morphology from the lost lectotype. 
That specimen (Schlegelmilch, 1976: pi. 28, fig. 3) is the original of 
Quenstedt, 1885, pi. 30, fig. 9, and represents a round-whorled, striate 
species of Polymorphites, which was described as P. lineatus (Quenstedt, 
1845) by Schlatter (1980: 86). 

The relationship between P. trivialis (Simpson, 1843) and P. 
polymorphus (Quenstedt, 1845) remains to be clarified: both have 
wide ranges of morphological variation, and they may be synonyms. 
P. trivialis is abundant in the lower half of the Brevispina Subzone, 
but two examples off! polymorphus (PI. 7, fig. 3) that are identical 
with the specimen figured by Dean et al. (1961: pi. 68, fig. 4) and 
Schlatter ( 1980: pi. 7, fig. 2) were found in bed 555 in the Jamesoni 
Subzone. They have broad whorls, widely spaced ribs and ventro- 
lateral tubercles that are characteristic of the most strongly 
ornamented forms of both species. 

Genus PLATYPLEUROCERAS Hyatt, 1867 

Platypleuroceras brevispina (J. de C. Sowerby, 1827) 

1827 Ammonites brevispina J. deC. Sowerby: 106, pi. 556, fig. 1. 

1843 Ammonites ripleyi Simpson: 11. 

1880/82 Aegoceras brevispina (J. de C. Sowerby); Wright: 361 

(1882), pi. 32, fig. 2, 3 (1880) (holotype, BM 43915, from 

Pabba, Inner Hebrides, Scotland). 
1909 Uptonia ripleyi (Simpson); Buckman: pi. 2 (holotype, WM 

106, probably from beds 544.7-546.4). 
1961 Platypleuroceras brevispina (J. de C. Sowerby); Dean et 

al.: pi. 69, fig. 1 (holotype). 

Range. From beds 544.6-550, Brevispina and Jamesoni Subzones; 
83 specimens. 

Platypleuroceras obsoleta (Simpson, 1843) 

1843 Ammonites obsoletus Simpson: 23. 

1882 Aegoceras brevispina (J. de C. Sowerby); Wright: 361, pi. 

50, figs 13, 14(BMC3126). 
1914 Uptonia obsoleta (Simpson); Buckman: pi. 92 (holotype, 

WM 157). 

Range. A single specimen in bed 544.7, Brevispina Subzone. 

Remarks. This is a single poorly preserved specimen with many 
ribs and ventro-lateral tubercles which is not good enough to elucidate 
the horizon of Simpson's larger and better preserved holotype. 

Platypleuroceras aureum (Simpson, 1855) PL 7, fig. 6 

1855 Ammonites aureus Simpson: 44 (non Ammonites aureus 

Young & Bird, 1822). 
71855 Ammonites tenuispina Simpson: 69 (the holotype is lost - 

seeHowarth, 1962: 111). 
1909 Platypleuroceras aureum (Simpson); Buckman: pi. 3 

(holotype, WM 107, from bed 546.2 or 546.5). 

Range. Beds 546. 1-546.5, Brevispina Subzone; 23 specimens. 



Remarks. P. aureum is a more evolute species than P. brevispina, 
and is bituberculate (ie. both umbilical and ventro-lateral tubercles are 
well developed). A small, typical specimen is figured in PL 7, fig. 6. 

Genus UPTONIA Buckman. 1897 

Uptonia jamesoni (J. de C. Sowerby, 1827) 

1827 Ammonites jamesoni J. de C. Sowerby: 105, pi. 555, fig. 1. 

71855 Ammonites ignotus Simpson, 1855: 61. 

71910 Uptonia ignota (Simpson); Buckman: pi. 21 (holotype, 

WM 159). 
1973 Uptonia jamesoni (J. de C. Sowerby); Donovan & Forsey: 

12, pi. 4, fig. 3 (neotype, BM C.40426, designated by 

Donovan & Forsey, 1973: 12, from Pabba, Inner Hebrides, 

Scotland). 

Range. Beds 550-560.3, Jamesoni Subzone; 23 specimens. 

REMARKS. Many of the 23 specimens in the Jamesoni Subzone are 
typical examples of the species, but they are mostly fragmentary, and 
none are preserved well enough to be figured. 

Uptonia lata (Quenstedt, 1845) 

1 845 Ammonites jamesoni J. de Sowerby, var. latus Quenstedt: 
88, pi. 4, fig. 1 (holotype, from Wiirttemberg, Germany). 

1980 Uptonia lata (Quenstedt); Schlatter: 1 13, pi. 12, figs 3, 4 
(from Wiirttemberg, Germany). 

Range. Beds 558 and 560.3, Jamesoni Subzone; 6 specimens. 

REMARKS. A more involute species than U. jamesoni, with much 
finer ribbing; identified according to Schlatter's (1980: 1 13) interpre- 
tation of Quenstedt's species. 

Genus TROPIDOCERAS Hyatt, 1867 

Tropidoceras futtereri Spath, 1923 PL 7, figs 10, 11 

1923a Tropidoceras futtereri Spath: 8. 

1928 Tropidoceras futtereri Spath; Spath: 228, pi. 16, fig. 8 

(holotype, BM C.23687, from bed 118b, Charmouth, 

Dorset). 

Range. Single specimens in beds 560.3 (top) and 568 (base), 
Masseanum Subzone. 

Tropidoceras masseanum (d'Orbigny), var. rotundum 
(Futterer, 1893) 

1 893 Cycloceras masseanum (d'Orbigny), var. rotundum Futterer: 
330, pi. 12, figs 3, 4 (holotype, from Wiirttemberg, Ger- 
many). 

1980 Tropidoceras masseanum (d'Orbigny) rotundum (Futterer); 
Schlatter: 138, pi. 19, fig. 4. pi. 20, figs 1, 2 (from 
Wiirttemberg, Germany). 

Range. Found only at the boundary of beds 567 and 568, Masseanum 
Subzone; 1 1 specimens. 

Genus PARINODICERAS Trueman, 1918 

Remarks . Although considered by Spath ( 1 93 8 : 8 1 ) to be a subgenus 
of Liparoceras, Parinodiceras (including its synonym Platynoticeras) 
is now thought to have been derived from Polymorphites and is there- 
fore placed in the family Polymorphitidae (Donovan, 1981: 111, 138). 



LOWER LIAS OF ROBIN HOOD'S BAY 



141 



Parinodiceras parinodum (Quenstedt, 1884) 

1884 Ammonites striatus parinodus Quenstedt: 225, pi. 28, fig. 

16. 
1938 Liparoceras (Parinodiceras) parinodum (Quenstedt); Spath: 

82, pi. 6, fig. 5 (from Radstock, Somerset), pi. 25, figs 1,4, 

5 (all from Wurttemberg. Germany). 
1976 Liparoceras (Parinodiceras) parinodum (Quenstedt); 

Schlegelmilch: 67, pi. 32, fig. 3 (lectotype, from Ofterdingen, 

Wurttemberg, Germany). 

RANGE. Beds 546.3, 548 and 554, Brevispina and Jamesoni Subzones; 
3 specimens. 



Family LIPAROCERATIDAE Hyatt, 1867 

Genus LIPAROCERAS Hyatt, 1867 

Subgenus LIPAROCERAS Hyatt, 1867 

Liparoceras (L.) cheltiense (Murchison, 1834) 

1834 Ammonites cheltiensis Murchison: 20, fig. 1. 

1904 Liparoceras cheltiense (Murchison); Buckman: pis 67, 67a 

(holotype, BM 74955a, from Gloucestershire). 
1938 Liparoceras cheltiense (Murchison); Spath: 46. 

Range. Two specimens in bed 562, Masseanum Subzone. 

Liparoceras (L.) heptangulare (Young & Bird, 1828) 

1828 Ammonites heptangularis Young & Bird: 263, pi. 14, fig. 1. 
1914 Liparoceras heptangulare (Young & Bird); Buckman: pis 

108 A-C (holotype, WM 170, probably from bed 575). 
1938 Liparoceras heptangulare (Young & Bird); Spath: 59. pi. 7, 

fig. 1 (BM C.2685, possibly from bed 575). 

RANGE. Single specimens in beds 57 1 , 575 and 577, Valdani Subzone; 
the best preserved is in bed 577. 

Liparoceras (L.) cf. naptonense Spath, 1938 

1938 Liparoceras naptonense Spath: 63, pi. 6, fig. 1 , pi. 9, fig. 7, 
pi. 10, fig. 6 (holotype, BM C. 1 2638, from Napton, Warwick- 
shire), pi. 14, fig. 6, pi. 16, fig. 10 (all from Warwickshire or 
Leicestershire). 

Range. Single specimens in beds 580 and 582.3, Luridum and 
Maculatum subzones. 

Liparoceras (L.) divaricosta (Trueman, 1919) PI. 8, fig. 1 

1919 Androgynoceras divaricosta Trueman: 278, pi. 22, fig. 1 
(holotype, BM C.38326, from Lincolnshire). 

1938 Liparoceras divaricosta (Trueman); Spath: 68, pi. 5, fig. 1 
(holotype). 

RANGE. A single specimen in bed 596.3, Figulinum Subzone. 



Genus AEGOCERAS Waagen, 1 869 
Subgenus AEGOCERAS Waagen, 1869 

Aegoceras (A.) maculatum (Young & Bird, 1822) 

PI. 7, figs 12, 13 

1822 Ammonites maculatus Young & Bird: 248, pi. 14, fig. 12. 
1828 Ammonites maculatus Young & Bird; Young & Bird: 259, 
pi. 14, fig. 9. 



1829 Ammonites arcigerens Phillips: 163, pi. 13, fig. 9. 
1835 Ammonites arcigerens Phillips; Phillips: 135, pi. 13, fig. 9. 
1875 Ammonites arcigerens Phillips; Phillips: 270, pi. 13, fig. 9. 
1 880/82 Aegoceras maculatum (Young & Bird); Wright: 368 ( 1 882), 

pi. 34 (1880), figs 1-3 (SM J18227). 4-7 (SM J18228). 
1912 Androgynoceras maculatum (Young & Bird); Buckman: 

pis 45 A, B (holotype, WM 493; from bed 590.61) 
1938 Androgynoceras maculatum (Young & Bird); Spath: 126, 

pi. 20, fig. 6 (BM C. 17752, from bed 590.61). 
1938 Androgynoceras maculatum (Young & Bird), var. rigida; 

Spath: 126, pi. 19, figs 2 (BM C.28175, from bed 590.61), 

13 (BM C.24601, from bed 590). 

1961 Androgynoceras maculatum (Young & Bird); Dean et al.\ 
pi. 70, fig. 4 (BM C. 17752; from bed 590.61 ) 

1962 Androgynoceras arcigerens (Phillips); Howarth: 112. pi. 
16, fig. 5 (holotype, BM 17139; from bed 590.61). 

1976 Androgynoceras (A.) maculatum (Young & Bird); 

Schlegelmilch: 162, pi. 33, fig. 9 (WM 493). 
1985 Androgynoceras (Beaniceras) luridum (Simpson); Phelps: 

350, pi. 1, fig. 3 (from bed 583). 
1985 Androgynoceras (Aegoceras) sparsicosta (Trueman); 

Phelps: 351, pi. 1, fig. 1 (from bed 585). 
1985 Androgynoceras (Aegoceras) maculatum (Young & Bird); 

Phelps: 350, pi. 2, fig. 8 (probably from bed 590.61). 

Range. Beds 581-590.63, Maculatum Subzone; 35 specimens. 

Remarks. Ammonites in bed 590.61 have a distinctive style of 
preservation, where the dark brown or black shell of the ammonite has 
small near-circular patches of white calcite; such a well-preserved 
example is figured in PI. 7, fig. 13. This type of preservation does not 
occur at any other level and allows the horizon of many of the figured 
specimens, including the holotype, to be identified precisely, as given in 
the list of figured specimens above. The lowest specimens in bed 58 1 are 
typical examples of the species, being much larger and more developed 
than is found in transitions from Aegoceras (Beaniceras) luridum. The 
lattertransitionsarerepresentedbytwospecimensinbed583.2determined 
as A. (A.) maculatum var. atavum (see discussion below). The specimen 
from bed 583 figured by Phelps as A. (Beaniceras) luridum is also an A. 
maculatum ; it has ribs on the venter that are bold and well developed and 
are not much reduced on a nearly flat venter as in B. luridum. 

Phelps (1985: 351) divided the Maculatum Subzone into a lower 
'Sparsicosta Zonule' and an upper 'Maculatum Zonule'. His lower 
division was based on the range of ammonites in the lower part of the 
Subzone that have low rib densities on their inner whorls of 16-18 
ribs per whorl, and were identified as Androgynoceras (Aegoceras) 
sparsicosta (Trueman). Such rib densities are not different from the 
rib densities of Aegoceras maculatum, and in any case sparsicosta is 
not an appropriate name for them. The real Androgynoceras 
sparsicosta (Trueman) (holotype figured Spath. 1938: pi. 5 fig. 7) is 
a species that develops swollen, quickly expanding whorls and has 
sharply bituberculate ribs from a diameter of about 25 mm. These 
features are typical of the genus Androgynoceras, and Phelps' (1985: 
pi. 1, fig. 1) 58 mm diameter ammonite from the lower part of the 
Maculatum Subzone does not show such features - it is a typical 
Aegoceras maculatum, as are all the specimens in Bairstow's collec- 
tion. Subdivision of the Maculatum Subzone on the basis of these 
ammonites is not followed here. 

Aegoceras (A.) maculatum (Young & Bird), var. atavum 
Spath, 1938 PI. 7, fig. 8 

1938 Androgynoceras maculatum (Young & Bird), var. atavum 
Spath: 127, pi. 20, fig. 3 (from Gloucestershire). 



142 



M.K. HOWARTH 




PLATE 8 

Fig. 1 Liparoceras (L.) divaricosta (Trueman). Bed 596.3. C. 39455, x 0.67: wholly septate. 
Fig. 2 Eoderoceras armatum (J. Sowerby ). Bed 499. near Bay Town, 0. 1 8 m above base of bed. CA 3885, x 1 . 

Fig. 3 Bifericeras donovani Dommergues & Meister. Bed 501 . 1 , near Bay Town. 0.22 m above base of bed. CA 3804; 3a, 3b, x 1 : 3c, d, x 2; wholly 
septate. 



LOWER LIAS OF ROBIN HOOD'S BAY 



143 



Range. Two specimens in bed 583.2, Maculatum Subzone. 

Remarks. This record is based on two specimens (C. 38873-74) 
from bed 583.2 that were determined by Spath ( 1938: 1 33) as belong- 
ing to his var. atavum; two other specimens in the same bed 
(C. 34887 1 -72) are typical of the normal variety of maculatum. C. 38873- 
74 are both only 34 mm diameter, possibly nearly adult, with reduced 
ribbing on the nearly flat venter (PI. 7, fig. 8). The whorl breadth is not 
as large as in Spath's (1938: 127, pi. 20, fig. 3) type of his variety, but 
otherwise they are closely similar. Spath ( 1 938: 1 28) himself remarked 
that an alternative place for such specimens might be in the genus 
Beaniceras, and it is possible that bed 583.2 might be the horizon from 
which the holotype of Aegoceras (Beaniceras) luridum was obtained 
(SM J3274, figured Dean et al, 1961: pi. 69, fig. 6). That holotype is 
somewhat larger (46 mm diameter), more complete and better pre- 
served than Bairstow's specimens, and no others that are as 
well-preserved have been found in beds 578-583. Spath's determina- 
tion as var. atavum for these Bairstow collection ammonites is retained 
here. 

Aegoceras (A.) maculatum (Young & Bird), var. leckenbyi 
Spath, 1938 



1938 



1938 



Androgynoceras maculatum (Young & Bird), var. leckenbyi; 
Spath: 126, pi. 13, fig. 2 (C.3741, from bed 590.1). 
Androgynoceras maculatum (Young & Bird), var. arcigerens 
(Phillips); Spath: 126, pi. 20, fig. 5 (from Dorset). 



Range. A single specimen in bed 590. 1 , Maculatum Subzone. 

Remarks. This variety is kept distinct from the normal type of A. 
maculatum only because it develops massive whorls with 'Liparoceras- 
type' of ornamentation at sizes of more than 100 mm diameter. 
Bairstow's specimen is 120 mm diameter, and a Dorset specimen with 
similar whorls at 120-150 mm diameter was figured by Spath ( 1938: 
pi. 20, fig. 5) as A. maculatum var. arcigerens (Phillips). 

Aegoceras (A.) lataecosta (J. de C. Sowerby, 1827) 

1 827 Ammonites lataecosta J. de C. Sowerby: 1 06, pi. 556, fig. 2. 
1880/82 Aegoceras lataecosta (J. de C. Sowerby); Wright: 365 

(1882), pi. 32, fig. 1 (1880) (holotype, BM 43916, from 

Drift, locality unknown). 
1938 Androgynoceras lataecosta (J. de C. Sowerby); Spath: 135, 

pi. 19, figs 4 (holotype, BM 43916), 6 (BM C.38562, from 

Staithes, Yorkshire). 

Range. Single specimens in beds 59 1 and 594, Capricomus Subzone. 
Aegoceras (A.) artigyrus (Brown, 1837) 



1837 
1843 
1855 
1884 
1889 
1938 



1973 



1973 



Ammonites artigyrus Brown: 26, pi. 19, fig. 5. 
Ammonites defossus Simpson: 15 
Ammonites defossus Simpson: 48 
Ammonites defossus Simpson: 78 
Ammonites artigyrus Brown: 20, pi. 19, fig. 5. 
Androgynoceras artigyrus (Brown); Spath: 158, pi. 14, fig. 
5, pi. 18, fig. 1, pi. 23, figs 3 (holotype, Manchester Museum 
LL.230, possibly from bed 593), 12. 14. 
Aegoceras maculatum (Young & Bird); Donovan & Forsey: 
14, pi. 4, fig. 1 (SM Bl 1945, paralectotype of Ammonites 
defossum Simpson, 1843). 

Aegoceras (Oistoceras) cf. figulinum (Simpson); Donovan 
& Forsey: 14, pi. 4, fig. 2 (SM B 1 1 946, lectotype of Ammo- 
nites defossum Simpson, 1843, designated by Donovan & 
Forsey). 



1985 Androgynoceras (Aegoceras) capricomus (Schlotheim), 
morphotype A. artigyrus; Phelps: 352, pi. 2, fig. 6. 

Range. Found by Bairstow only in bed 593, Capricomus Subzone, 
4 specimens, but Phelps' figured specimen is from bed 595.2. 

Remarks. Donovan & Forsey's (1973: 14) designation of the Robin 
Hood's Bay specimen SM B 1 1946 as lectotype of Ammonites defossus 
Simpson, 1843, has consequences for the position of both the genus 
Defossiceras Buckman, 1913, and Simpson's species defossus. That 
lectotype and the paralectotype (SM B 1 1945), also figured for the first 
time by Donovan & Forsey, are closely similar to each other, and both 
have robust, quickly expanding whorls and moderately fine ribs on the 
inner whorls. The robust whorls are not like the slender whorls of 
Oistoceras at similar sizes, and the ribs are more closely spaced on the 
inner whorls than in Aegoceras (A. ) maculatum. The ribs are projected 
slightly forwards on the venter of both lectotype and paralectotype of 
defossus, but a varying amount of projection of the ribs on the venter 
occurs in Capricomus Subzone species of Aegoceras (A.), though it is 
never as pronounced as in the later subgenus Oistoceras. The holotype 
of A. (A. ) artigyrus figured by Spath (1938) has similarly robust whorls, 
the same rib-density on the inner whorls and similar slight forward 
projection of the ribs on the venter. This is considered here to be the best 
place for defossus, and places the genus Defossiceras as a junior 
synonym of Aegoceras (A.) (ie. not a synonym of the subgenus 
Oistoceras), and the species defossus as ajunior synonym of Aegoceras 
(A.) artigyrus. 

Subgenus BEANICERAS Buckman, 1913 

Aegoceras (Beaniceras) luridum (Simpson, 1855) 

1855 Ammonites luridus Simpson: 46. 

1913 Beaniceras luridum (Simpson); Buckman: pi. 73 (holotype, 
SM J3274). 

1961 Beaniceras luridum (Simpson); Dean et al.: pi. 69, fig. 6 
(holotype, SM J3274, possibly from bed 583.2) (see de- 
scription of Maculatum Subzone, p. 150 below). 

Range. Beds 578. 1-580, Luridum Subzone; 8 specimens; possibly 
also from bed 583.2, Maculatum Subzone. 

Remarks. Although the eight specimens in beds 578 and 580 are 
crushed, they are much larger (up to 37 mm diameter) and more com- 
pressed than A. (B.) centaurus (d'Orbigny ), and so can be confidently 
referred to A. (B.) luridum. 

Subgenus OISTOCERAS Buckman, 1911 

Aegoceras (Oistoceras) sinuosiforme Spath, 1938 

PI. 7, fig. 14 

1938 Oistoceras sinuosiforme Spath: 167, pi. 1 8, fig. 6, pi. 19, fig. 
7 (holotype, BM C.38564), pi. 26, figs 6, 7, 9 (all Spath's 
figured specimens are from Lincolnshire). 

Range. Beds 596.2-598. 1 , Figulinum Subzone; 93 specimens. 

REMARKS. A.(0.) sinuosiforme has more widely spaced ribs and 
much less well-developed chevrons on the venter than A. (O. ) figulinum. 

Aegoceras (Oistoceras) angulatum (Quenstedt, 1856) 

1856 Ammonites maculatus angulatus Quenstedt: 121, pi. 14, fig. 
12. 

1885 Ammonites maculatus angulatus Quenstedt; Quenstedt: 270, 
pi. 34, fig. 11. 



144 
1938 



1976 



1985 



Oistoceras angulation (Quenstedt); Spath: 1 7 1 , pi. 2 1 , fig. 5 
(from France), pi. 22, fig. 5 (from Lincolnshire), pi. 26, figs 
10, 12 (both from Lincolnshire). 

Androgynoceras (Oistoceras) angulatum (Frebold, 1922) 
(sic); Schlegelmilch: 69, pi. 34, fig. 4 (from Germany). 
Androgynoceras (Oistoceras) angulatum (Quensedt); 
Phelps: 354, pi. 2, fig. 4 (from Germany). 

Range. Bed 599, Figulinum Subzone; 1 specimen. 

Remarks. Quenstedt ( 1 856: 1 2 1 ) had specimens from Metzingen 
and Iggingen, Germany, and he figured one from the former locality. So 
the original of Quenstedt (1884: pi. 34, fig. 11), refigured by both 
Schlegelmilch (1976: pi. 34, fig. 1 1 ) and Phelps (1985: pi. 2, fig. 4), is 
almost certainly a syntype and can be designated lectotype: its designa- 
tion by Schlegelmilch as the neotype is not correct. A. (O.) angulatum 
is more evolute, has more slowly expanding whorls, has no ventro- 
lateral tubercles, and has fewer ribs on the inner whorls, than A. (O.) 
figulinum. The angle of the ribbing varies between rectiradiate and 
prorsiradiate in both species. 

Aegoceras (Oistoceras) figulinum (Simpson. 1855) 

1855 Ammonites figulinus Simpson: 47. 
1855 Ammonites omissus Simpson: 47. 
1876 Aegoceras defossum (Simpson); Blake: 282, pi. 8, fig. 9 

(SM J17988. see Donovan & Forsey, 1973: 13). 
1911 Oistoceras figulinum (Simpson): Buckman: pi. 26A 

(holotype, WM 115). 
191 1 Oistoceras omission (Simpson); Buckman: pi. 27 (holotype, 

WM 502, now lost). 
1938 Oistoceras figulinum (Simpson); Spath: 162, pi. 19, fig. 10 

(BM C. 17988), pi. 22, fig. 8 (BM 37973a). 
1938 Oistoceras omissum (Simpson): Spath: 170. pi. 21. fig. 3 

(BM 38561). 
1955 Oistoceras aff. figulinum (Simpson); Howarth: 1 6 1 , pi. 11. 

fig. 4 (SM J35968. from bed 600.4). 
1976 Androgynoceras (Oistoceras) figulinum (Simpson): 

Schlegelmilch: 69, pi. 34, fig. 3 (WM 115. holotype). 
1985 Androgynoceras (Oistoceras) figulinum (Simpson): Phelps: 

353, pi. 2, fig. 1 (from bed 600.2). 
1987 Oistoceras figulinum (Simpson); Dommergues: pi. 1 1, figs 

5.6. 

Range. Beds 600.2 and 600.4, Figulinum Subzone; 20 specimens. 

Remarks. This is the most highly developed species of Oistoceras. 
which has fine ribs on the inner whorls, small ventro-lateral tubercles, 
and well-developed chevrons in the ribs that are connected together 
into a rudimentary pseudo-keel along the middle of the venter. 

Other species of Oistoceras from Yorkshire: 

1. A. (O. ) cunicome (Schloenbach, 1863);Spath. 1938: 164, pi. 19, 
fig. 1 1 (BM C. 19228; indeterminate inner whorls), pi. 22, fig. 9 
(BM C.6235); both of Spath's figured specimens were probably 
from Staithes. not Robin Hood's Bay. 

2. ?A. (O.) anguliferum (Phillips, 1829: 163, pi. 13, fig. 19: 1835: 
135, pi. 13, fig. 19; 1875: 270, pi. 13, fig. 19); the type specimen 
is lost, and Phillips' figure cannot be interpreted. 

Genus ANDROGYNOCERAS Hyatt, 1867 

Androgynoceras heterogenes (Young & Bird. 1828) 
1828 Ammonites heterogenes Young & Bird: 264, pi. 14, fig. 7. 



M.K. HOWARTH 

1880/82 Aegoceras heterogenum (Young & Bird): Wright: 370 

(1882), pi. 35, figs 4-6, (1880) (SMJ18229), pi. 36, figs 1- 

4 (1880) (BMC. 1870). 
1912 Androgynoceras heterogenes (Young & Bird): Buckman: 

pi. 46 (holotype, WM 195). 
1938 Androgynoceras heterogenes (Young & Bird); Spath: 1 13, 

pi. 13, figs 7a (BM C.19225). 7b (BM C.38457). pi. 20, fig. 

2 (BM C. 38496. as van gigas, from bed 590.61 ). 

Range. Maculatum Subzone: Bairstow found single specimens in 
beds 583.2 and 588, but BM C.38496 definitely came from bed 590.6 1 . 
and the other figured specimens are probably from bed 590.63. 



Order NAUTILOIDEA 

Family NAUTILIDAE d'Orbigny, 1840 
Genus CENOCERAS Hyatt. 1883 

Cenoceras striatus (J. Sowerby, 1817) 

1817 Nautilus striatus J. Sowerby: 183. pi. 182 (3 figures, all 

syntypes. from Dorset). 
1829 Ammonites annularis Phillips: 163, pi. 12, fig. 18: 1835: 

134, pi. 12, fig. 18; 1875: 263, pi. 12, fig. 8. 
1855 Ammonites heterogeneus Simpson: 33. 
1956 Cenoceras striatus (}. Sowerby); Kummel: 362, pi. 3, figs 1, 

2 (BM 43852, from Dorset). 
1962 Cenoceras heterogeneum (Simpson); Howarth: 96. pi. 13, 

fig. 1 (holotype. WM 442). 
1962 Cenoceras annulare (Phillips): Howarth: 96, pi. 13, fig. 2 

(holotype, WM 62). 

Range. Bairstow found single specimens in beds 464.32, 468 (both 
Simpsoni Subzone) and 505.1 (Taylori Subzone). 



BIOSTRATIGRAPHY 



In the description below the ammonite distribution and the place- 
ment of the boundaries are discussed for all the zones and subzones 
in Robin Hood's Bay. Additionally, it is noteworthy that Wine Haven 
at the south-eastern end of the bay has recently been proposed as the 
world standard for the definition of the base of the Pliensbachian 
Stage. 

The scheme of ammonite zones used here is based on that regular- 
ized by Dean. Donovan & Howarth (1961), with a few later 
refinements to the details of some of the definitions. Cariou & 
Hantzpergue (1997) used the same scheme of divisions for the 
Sinemurian and Lower Pliensbachian in eastern France and the 
central Mediterranean area. The distribution of the ammonites, on 
which the biostratigraphical divisions are based, is shown in detail in 
Figs 2 1 , 22, 24 and 25, which give the number of specimens of each 
species found in each bed. and a visual indication of the range of each 
species. 

LOWER SINEMURIAN 

SEMICOSTATUM ZONE, Sauzeanum Subzone, beds 418- 
429.64. No ammonites were found in beds 418-420. which are the 
first 2.48 m of strata exposed above the lowest level to which the tide 
ever falls in Robin Hood's Bay. Above this, Euagassiceras occurs up 
to about the middle of the subzone. and Coroniceras (Arietites) 
alcinoe occurs in a broad middle part of the subzone; both are 



LOWER LIAS OF ROBIN HOOD'S BAY 



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Fig. 21 Distribution of ammonites in the Lower Sinemurian of Robin 
Hood's Bay. 

characteristic of the Sauzeanum Subzone. Amioceras semicostatum 
is common through most of the subzone. 

TURNERI ZONE, Brooki Subzone, beds 429.7-433.2. The only 
ammonites found in the beds that are allocated to this subzone are 
seven Caenisites brooki in the middle part and two Amioceras sp. 
indet. in the middle and lower beds. Caenisites brooki probably only 
occurs in the upper or top part of the subzone (Dean et al, 1961: 
453), and the ammonite Caenisites preplotti Spath, which is charac- 
teristic of the base of the subzone, does not occur in Robin Hood's 
Bay. So the Brooki Subzone has to be defined according to the 
boundaries of the adjoining subzones: the highest occurring 
Coroniceras (Arietites) alcinoe in bed 429.64 at the top of the 
Sauzeanum Subzone defines the base of the Brooki Subzone at the 
bottom of bed 429.7, and the appearance of Microderoceras birchi in 
bed 433.3 at the base of the Birchi Subzone defines the top of the 
Brooki Subzone at the top of bed 433.2. 

Birchi Subzone, beds 433.3^146.2. This subzone is generally con- 
sidered to correspond to the range of Microderoceras: five M. birchi 
occur in bed 433.3, so defining the base of the subzone, and a single 



M. scoresbyi occurs in bed 441.2 at the middle of the subzone. The 
top of the subzone is delimited by the appearance of the first 
Asteroceras at the base of the Obtusum Zone. Caenisites brooki 
persists into the basal bed (433.3) of the Birchi Subzone, and the 
same bed also contains 24 examples of Caenisites turneri. 
Promicroceras capricornoides appears just above the lowest part 
and extends to the top of the subzone. There are no other ammonites 
in the subzone. 

UPPER SINEMURIAN 

OBTUSUM ZONE, Obtusum Subzone, beds 446.31-446.5. The 
base of both zone and subzone is drawn at the first appearance of a 
single Aste roceras in bed 446.3 1 ; that specimen is a definite example 
of the genus, but is not specifically determinable. The only specimen 
of A. obtusum that was found occurs in the overlying bed 446.32, and 
A. confusum is more common in beds 446.32 and 446.33. 
Promicroceras capricornoides persists into the lowest two beds of 
the Obtusum Subzone, then is immediately replaced by P. planicosta 
for the remainder of the subzone: the two species do not overlap. 
Other ammonites are Xipheroceras dudressieri (confined to the 
subzone) and X. zipluts, Epophioceras landrioti in the upper half and 
Cymbites laevigatas at the top of the subzone. 

Stellare Subzone, beds 447-455.1. The base of the subzone is 
placed at the first appearance of the distinctive index species 
Asteroceras stellare. which ranges up to the middle of the subzone, 
and the top is limited by the first Eparietites at the base of the 
Denotatus subzone. In the upper half of the subzone the index 
species is replaced by Asteroceras blakei, which persists into the 
overlying subzone. Aegasteroceras crassum appears at about the 
middle of the subzone, then A. sagittarium occurs in the top part. 
Promicroceras planicosta is very common in all but the highest beds 
of the subzone, and 262 specimens were collected by Bairstow. 
Other ammonites in the subzone are Cymbites laevigatus, Xiphero- 
ceras ziphus, and Epophioceras landrioti near the base. 

Denotatus Subzone, beds 455.2—162. The base of this subzone is 
placed at the first appearance of the genus Eparietites, ie. the new 
species E. bairstowi, which is more evolute and has thicker and more 
massive whorls than any other Eparietites. The main species ranging 
through the middle and upper parts and up into the Simpsoni Subzone 
is E. impendens. From the subzone below Asteroceras blakei, 
Aegasteroceras crassum and A. sagittarium persist into the bottom 
and middle parts of the Denotatus Subzone. Cymbites laevigatus 
occurs throughout the subzone, and the Schlotheimid Angulaticeras 
sp. indet. occurs in the top two beds. 

OXYNOTUM ZONE, Simpsoni Subzone, beds 463-171. The 
base of the subzone is placed at the first appearance of the index 
species Oxynoticeras simpsoni in bed 363. where there are two large 
specimens that show typical characters of the species; there are four 
more specimens in bed 464.3, poorly preserved examples in beds 
465 and 466, then the species becomes common in bed 467 and 468 
in the mid to upper part of the subzone. 

From the subzone below, Eparietites impendens persists into beds 
463—164.32, where it overlaps with O. simpsoni in the bottom 1 .68 m 
of the Simpsoni Subzone. In fact at its highest level in bed 464.32 
there are many typical E. impendens. A similar overlap between E. 
impendens and O. simpsoni is also found in the top part of the 
Frodingham Ironstone near Scunthorpe, Lincolnshire. 

Gagaticeras is characteristic of the upper half of the Simpsoni 
Subzone. from bed 467 upwards, where there are many specimens 
belonging to four species. Palaeoechioceras occurs in bed 467, and 



146 



M.K. HOWARTH 



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Fig. 22 Distribution of ammonites in the Upper Sinemurian of Robin Hood's Bay. 



LOWER LIAS OF ROBIN HOOD'S BAY 



147 



Cymbites laevigatus and Angulaticeras sp. indet. occur in the lower 
half of the subzone. 

Oxynotum Subzone, beds 472.1-486.2. More involute and com- 
pressed Oxynoticeras like O. oxynotum rather than O. simpsoni first 
occur in bed 472. 1 , so the base of the subzone is placed at that level. 
Better specimens occur higher in the subzone, as well as fragments 
of large specimens. Other oxynoticeratids present are two possible 
specimens of Paroxy notice ras salisburgense in the lower half, and 
Gleviceras doris and G. guibalianum in the upper half of the subzone. 
Angulaticeras sp. indet., Bifericeras bifer and B. cf. vitreum, also 
occur in the upper half of the subzone. 

RARICOSTATUM ZONE, Densinodulum Subzone, beds 486.3 
and 487. The base of the subzone is placed at the first appearance of 
Crucilobiceras in bed 486.3 and the top is limited by the first 
occurrence of Echioceras in bed 488 marking the base of the 
Raricostatoides Subzone. So the Densinodulum Subzone consists 
only of the 1.0 m thick beds 486.3 and 487. C. densinodulum is 
abundant in bed 486.3, but there are no other ammonites in the 
subzone. 

Raricostatoides Subzone, beds 488^493.5. The base of the subzone 
is placed at the first appearance of Echioceras: E. raricostatoides in 
the basal one-third of the subzone is followed by E. intermedium in 
the middle part, then by Paltechioceras planum in the upper one- 
third of the subzone. Crucilobiceras densinodulum persists from the 
subzone below into the basal bed, and the only other ammonite in the 
subzone is Eoderoceras hastatum in the upper part. 

Macdonnelli Subzone, beds 494-495.7. This subzone is based on 
the range of the index species Leptechioceras macdonnelli, which 
occurs in the top and bottom beds and does not range higher. The 
earliest Eoderoceras armatum occurs in the bottom bed, and the first 
Polymorphitid, Gemmellaroceras tubellum, occurs in the top bed. 
The only other ammonites present are the Oxynoticeratids Gleviceras 
guibalianum near the top of the subzone and Radstockiceras 
buvignieri in the bottom bed. The latter record seems to be the first 
provable occurrence of Radstockiceras in the Raricostatum Zone. 

Aplanatum Subzone, beds 496-500. The base of the subzone is 
placed at the first occurrence of Paltechioceras regustatum, and P. 
tardecrescens (of which P. aplanatum is a synonym) becomes abun- 
dant in the middle and upper parts of the subzone. Paltechioceras 
first occurs in the top part of the Raricostatoides Subzone, but the 
genus is much more common in the Aplanatum Subzone and does 
not range higher. Another ammonite that is characteristic of the 
Aplanatum Subzone is Eoderoceras armatum, which first appears as 
rare examples in the Macdonnelli Subzone, but becomes much more 
common in the Aplanatum Subzone; it ranges up to 0. 15 m below the 
top of bed 499. but it does not occur higher and does not overlap with 
Apoderoceras in the Taylori Subzone. Gemmellaroceras tubellum is 
common in the middle part, and Gleviceras guibalianum occurs in 
the lower part of the subzone. 

LOWER PLIENSBACHIAN 

The exposures at the base of the cliff in Wine Haven, Robin Hood's 
Bay. have recently been proposed as the Global Stratotype Section 
and Point (GSSP) for the base of the Pliensbachian Stage (Hesselbo 
et al, 2000). The sequence across the Sinemurian/Pliensbachian 
boundary is sufficiently expanded and rich in ammonites here to be 
suitable for such an important global reference section. Hesselbo et 
al.'s (2000: 604, fig. 4) stratigraphical sequence is closely similar to 
the sequence described here, as are their ammonite records and 
identifications. Their bed 73 at the base of the Pliensbachian is the 



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1 2 Gemmellaroceras tubellum 



2 Eoderoceras armatum 
- 2 Gleviceras guibalianum 
2 Gemmellaroceras tubellum 



Fig. 23 The distribution of ammonites close to the Sinemurian/ 
Pliensbachian boundary in Robin Hood's Bay. The ammonites listed 
include those collected from the exposures across the boundary at both 
Wine Haven and the foreshore immediately east of Robin Hood's Bay 
town; there are no significant differences in the distribution of 
ammonites at the two exposures. 

same as bed 501 here (see the correlation table of Fig. 19), and their 
photograph (Hesselbo et al. 2000: fig. 3) shows the nodules of their 
bed 72 (=bed 500 here) and the basal reference point of the 
Pliensbachian low in the cliff at Wine Haven. 

Fig. 23 shows details of the stratigraphical distribution of 
Bairstow's ammonites at the Sinemurian/Pliensbachian boundary. 
The first ammonites to occur above the boundary are 16 Bifericeras 
donovani Dommergues & Meister (one is figured in PI. 8. fig. 3) and 



148 



M.K. HOWARTH 



two Apoderoceras subtriangulare (Young & Bird) (PI. 5, fig. 8) 
0.13-0.22 m above the bottom of bed 501.1. Hesselbo etal. (2000) 
did not find ammonites in the 1.8 m of strata below the base of the 
Pliensbachian (ie. in the nodules of bed 500 and the shales of bed 
499). Bairstow did not find ammonites in bed 500, but he collected 
six specimens of Eoderoceras armatum from bed 499 in the middle 
of the bay near Robin Hood's Bay town. These are well-preserved, 
readily identifiable examples of the species (one is figured in PI. 8, 
fig. 2), and Bairstow's records show that they were collected 0.15- 
0.37 m above the base of that bed. A single E. armatum, less 
well-preserved than those lower down, but still readily identifiable 
with that species, was collected from Wine Haven high in bed 499, 
only 0.15 m from the top. The thickness of strata across the 



Sinemurian/Pliensbachian boundary from which no ammonites have 
been collected is thus reduced to only 0.36 m. 

JAMESONI ZONE, Taylori Subzone, beds 501.1-537. The base 
of the subzone (and the Jamesoni Zone, and the Pliensbachian Stage) 
is placed at the bottom of bed 501.1, which contains the lowest 
occurrence of the characteristic genus Apoderoceras, and this is the 
only horizon at which Bifericeras donovani Dommergues & Meister 
occurs. Phricodoceras taylori and other species of Phrkodoceras 
are present through much of the subzone and do not range higher. 
Many examples of Gemmellaroceras tubellum are found near the 
base of the subzone, and the larger species G. rutilans occurs in the 
top part. A single specimen of Radstockiceras buvignieri was also 



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Fig. 24 Distribution of ammonites in the Jamesoni Zone, Lower Pliensbachian, of Robin Hood's Bay. 



LOWER LIAS OF ROBIN HOOD'S BAY 



149 



found in the subzone, along with widely scattered specimens of 
Tragophylloceras numismale. 

Polymorphus Subzone, beds 538-544.5. The first appearance of 
Polymorphites marks the base of this subzone. The earliest species is 
P. caprarius which occurs in the bottom half of the subzone, fol- 
lowed by P. trivialis in the upper half. The latter species extends into 
the Brevispina Subzone, and other species occur in the Jamesoni 
Subzone. The only other ammonites in this subzone in Robin Hood's 
Bay are the last examples of Tragophylloceras numismale, a single 



Hype rde roc eras sp. indet. low in the subzone and the oxynoticeratid 
genus Radstockiceras: the small species R. sphenonotum is confined 
to the Polymorphus Subzone, and another much larger fragment of a 
Radstockiceras was found in bed 544.4 near the top of the subzone. 

Brevispina Subzone, beds 544.6-549. The base is placed at the first 
appearance of Platypleuroceras brevispina, which is common 
throughout the subzone and extends up to its highest occurrence in 
the basal bed of the Jamesoni Subzone. Many of those in beds 544 
and 546 are large, crushed and fragmentary, though they show the 



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Fig. 25 Distribution of ammonites in the Ibex and Davoei Zones, Lower Pliensbachian. and in the base of the Margaritatus Zone, Upper Pliensbachian, of 
Robin Hood's Bay. 



150 



M.K. HOWARTH 



typical features of the species. The more evolute P. aureum occurs in 
the middle part of the subzone, and the more finely ribbed P. obsoleta 
is represented by only one specimen in Bairstow's collection from 
the lower part of the subzone. Polymorphites trivialis is common in 
the lower and middle parts of the subzone, and there are rare 
occurrences of Radstockiceras, Parinodiceras and Tragophylloceras. 

Jamesoni Subzone, beds 550-560.3 (except the top 0.08m). The 
base of the Jamesoni Subzone is placed at the first occurrence of 
Uptonia jamesoni in bed 550, and the index species then extends 
through the full thickness of the subzone, to which it is confined. The 
more finely ribbed U. lata occurs less commonly in the upper half of 
the subzone, and Polymorphites bronni is also characteristic of the 
upper half of the subzone. The only other ammonites in the subzone 
are the highest occurring specimens of Platypleuroceras brevispina 
in bed 550. Polymorphites polymorphus at the middle of the subzone, 
and rare examples of Parinodiceras and Tragophylloceras. 

IBEX ZONE, Masseanum and Valdani Subzones, beds 560.3 (top 
0.08 m)-577. These two subzones are delimited according to the 
distribution of species of Tropidoceras and Acanthopleuroceras: 
Tropidoceras occurs in both subzones and Acanthopleuroceras only 
in the Valdani Subzone. Unfortunately ammonites are rare in this 
interval in Robin Hood's Bay, and the few examples of these genera 
are not well-preserved. None, however, have the definite bituberculate 
ribs of Acanthopleuroceras, so they all have to be identified as 
Tropidoceras, in which umbilical tubercles are much reduced or 
absent. The lowest example (PI. 7. fig. 10) from the top 0.08 m of bed 
560.3 and another (PI. 7, fig. 1 1 ) from near the bottom of bed 568 are 
best determined as T futtereri (Spath), while several specimens from 
the boundary of beds 567 and 568 have the much more massive ribs 
at larger sizes and the more widely spaced ribs of T. masseanum 
(d'Orbigny), var. rotundum (Futterer). The base of the Masseanum 
Subzone (and of the Ibex Zone) is placed 0.08 m below the top of bed 
560.3 to include this earliest T. futtereri, and bed 568 probably 
belongs to the same subzone. 

In the absence of Acanthopleuroceras there is no good evidence 
for the position of the base of the Valdani Subzone, so it is placed 
provisionally at the bottom of bed 571 from the occurrence of 
Liparoceras (L.) heptangulare. There are two more specimens of 
that species in beds 575 and 577. According to Spath ( 1938: 59) L. 
(L.) heptangulare might be confined to the Valdani Subzone (ie. 
Spath's 'Centaurus Subzone' ), so its presence in beds 57 1-577 (7.66 
m thick) suggests that they are probably of Valdani Subzone age. 

The only other ammonites in either subzone are two Liparoceras 
(L.) cheltiense low in the Masseanum Subzone, one Tragophylloceras 
loscombi high in the same subzone, and Lytoceras fimbriatum in the 
upper part of the Masseanum Subzone and throughout the Valdani 
Subzone. The latter species becomes more common in the Luridum 
Subzone. 

Luridum Subzone, beds 578. 1-580. The presence of eight Aegoceras 
(Beaniceras) luridum in beds 578.1, 578.5 and 580 is sufficient 
evidence to refer beds 578 to 580 to the Luridum Subzone. Other 
ammonites in this subzone are a single Liparoceras (L.) cf. 
naptonense, two Liparoceras (L.) sp. indet. and 14 examples of 
Lytoceras fimbriatum. 

DAVOEI ZONE, Maculatum Subzone, beds 58 1-590.7. The base 
of this zone and subzone is placed at the bottom of bed 581 which 
contains the lowest Aegoceras (A.) maculatum (PI. 7, fig. 12). Two 
more, typical, examples occur in bed 582.3, then there are many 
well-preserved specimens at higher levels, especially in beds 590.6 1 
and 590.63. Other ammonites in the Maculatum Subzone are A. (A.) 
maculatum vars atavum and leckenbyi. Liparoceras (L.) cf. 



naptonense, Androgynoceras heterogenes and Lytoceras sp. indet. 
See remarks on the identification of Aegoceras maculatum (p. 141) 
for discussion of the division of the Maculatum Subzone into smaller 
units. 

Capricornus Subzone, beds 59 1-596. 1 . The base of the subzone is 
placed at earliest occurrence of Aegoceras (A. ) lataecosta in bed 59 1 . 
This is 1.83 m above the highest A. (A.)maculatum in bed 590.63, but 
the intervening strata (beds 590.64-590.7) did not yield any ammo- 
nites and are retained in the Maculatum Subzone. The only other 
ammonites in the subzone are A. (A.) artigyrus, which has more 
massive whorls and coarser ribbing than lataecosta, and a number of 
poorly preserved Aegoceras (A.) sp. indet. 

Figulinum Subzone, beds 596.2-600.5. This subzone is based on 
the range of the subgenus Oistoceras. The index species, Aegoceras 
(Oistoceras) figulinum, occurs in beds 600.2 and 600.4 near the top 
of the subzone, but the base of the subzone is placed at the lowest 
appearance of A. (O.) sinuosiforme in bed 596.2. This and A. (O.) 
angulation in bed 600.2 have more widely spaced ribs thanfigulinum, 
especially on the inner whorls. The only other ammonite in the 
subzone is a single Liparoceras (L.) divaricosta in bed 596.3 (PI. 8, 

fig. 1). 

The top of the subzone is limited by the base of the Stokesi 
Subzone ( Margaritatus Zone, Upper Pliensbachian ), which is placed 
at the first appearance of Amaltheus stokesi in bed 600.6. There are 
other examples of A. stokesi in bed 600.8 and at higher levels in the 
Stokesi Subzone. Aegoceras (Oistoceras) figulinum and Amaltheus 
stokesi are confined to their respective subzones in Robin Hood's 
Bay and their ranges do not overlap. 



Acknowledgements. I wish to thank the late Leslie Bairstow for en- 
trusting me with his unfinished manuscripts in the hope that 1 would be able 
to complete them in a form suitable for publication. Mrs L.M. Spencer kindly 
helped me to retrieve those manuscripts when he left his London home in 
1985. Thanks are also due to Mr. Peter Jones of King's College, Cambridge, 
for information about Bairstow during his time at that college, to Rosalind 
Moad for allowing me to examine the original copy of Bairstow's 1930 
Fellowship Dissertation, which is held in the Archive Centre at King's 
College, and to Professor D.T. Donovan for reading the manuscript and 
suggesting several significant improvements. 



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revision of the English Wealden Flora, III: Czekanowskiales, 
Ginkgoales & allied Coniferales. 2001. Pp. 1-82. £43.40 

No. 2 The Cenozoic Brachiopod Terebratula: its type species, 

neotype, and other included species — Gough's Cave 1 
(Somerset, England): a study of the pectoral girdle and upper 
limbs — Systematic affinity of Acroporella assurbanipali Elliott 
(Dasycladaceae), with notes on the genus Neomeris Palyn- 
ological zonation of Mid-Palaeozoic sequences from the 
Cantabrian Mountains. NW Spain: implications for inter- 
regional and interfacies correlation of the Ludford/Pridoli and 
Silurian/Devonian boundaries, and plant dispersal patterns. 

2001. Pp. 83-162. £43.40 

Volume 58 

No. 1 Gough's Cave 1 (Somerset, England): a study of the axial 

skeleton — Upper Ordovician brachiopods from the Anderken 
Formation, Kazakhstan: their ecology and systematics. 

2002. Pp. 1-80. £43.40 



CONTENTS 

81 The Lower Lias of Robin Hood's Bay, Yorkshire, and the work of Leslie Bairstow 

M.K. Howarth 




Bulletin of The Natural History Museum 
GEOLOGY SERIES 

Vol. 58, No. 2, November 2002 



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Issued 26 June 2003 



Gough's Cave 1 (Somerset, England): a stuidjsroiw museum 



of the pelvis and lower limbs 



PRgSFMTPp 



ERIKTRINKAUS 

Department of Anthropology, Campus Box 1114, Washington University, St. Louis, MO 63 130, USA 



PALAEONTOLOGY UBRARY J 



SYNOPSIS. The lower limb remains of Gough's Cave 1 retain most of the pelvis, both femora, one complete tibia and portions 
of the other, sections of both fibulae, two tarsals and three metatarsals. They are those of a largely average European Mesolithic 
young adult male. Overall diaphyseal robusticity is generally similar to that of other Mesolithic specimens, even though the fibula 
and third metatarsal appear gracile. Musculo-ligamentous attachment areas are generally weakly marked. The proximal femora 
and the femoral diaphyses exhibit a clear asymmetry, especially in their neck-shaft angles and diaphyseal dimensions, which is 
is accompanied in the pelvis by a greater degree of left iliac lateral flare. These aspects are associated with a pelvis that combines 
several distinctly male characteristics with an overall pelvic aperture shape which is female. 



INTRODUCTION 



The Gough's Cave 1 skeleton retains a largely complete pelvis 
(which has been permanently articulated), both femora very well 
preserved, most of the right tibia and fibula, portions of the left tibia 
and fibula, the complete right talus and cuboid, and three complete 
metatarsals. As such, Gough's Cave 1 retains essentially complete 
anatomy on at least one side from the L5-S 1 articulation to the talo- 
calcaneal articulation, with additional data from the subtalar skeleton. 



MATERIALS 



The description of the Gough's Cave 1 lower limb remains includes 
extensive osteometries (Tables 1-5, 10, 11, 16-21). To evaluate 
some of these dimensions and the resultant proportions, comparative 
summary statistics (as available) are included for other European 
Mesolithic remains. These include remains from the sites of Arene 
Candide, Los Azules, Bichon, Birsmatten, Bottendorf, Riparo 
Continenza, Culoz, Gramat, Grotte des Enfants, Hoedic. Holmegard, 
Koelbjerg, Kosor Glas, Loschbour, Moita do Sebastiao, Molara, 
Mondeval, Muge (N - <57), Obercassel, Parabita, Le Peyrat, Le 
Rastel, Rochereil, Romanelli, Romito, Riparo Tagliente, San Teodoro, 
Sejr0, Teviec, Unseburg, Uzzo, Vaegens0, Vatte di Zambana, and 
Veryier 1 (Pittard & Sauter, 1946;Graziosi, 1947;Combier&Genet- 
Varcin, 1959; Barral & Primard, 1962; Genet- Varcin et at, 1963; 
Patte, 1968; Cremonesi et ai, 1972; Ferembach, 1976; Paoli et al, 
1980; Holliday, 1995; Holt, 1999; Churchill, pers. comm.). These 
comparative remains vary in age from terminal Paleolithic to well 
within the western European Mesolithic, approximately between 
1 2,000 and 6,000 years B. P. They should bracket reasonably well the 
Gough's Cave 1 remains in age. 

The most detailed metrics are available for the Gramat, Hoedic, 
Rochereil and Teviec remains, but the other specimens fill out the 
samples for the more commonly reported measurements (e.g., long 
bone lengths and diaphyseal diameters). For diaphyseal metrics, the 
femoral (proximal and midshaft), tibial (proximal) and fibular 
(midshaft) samples are dominated by the large sample from Muge. 
Consequently, when the Muge sample is significantly different from 
the remainder of this 'Mesolithic' sample, summary statistics for it 
are provided in addition to those for the total sample. 



Of the 39 comparative specimens other than those from Muge, 26 
are male, 12 are female and 1 has unknown sex. In the Muge femoral 
sample (the largest sample for the bones providing relevant data), 33 
are male and 24 are female. This is therefore a male biased sample, 
but given the probable male sex of Gough's Cave 1, this is not 
inappropriate. 



METHODS 

The majority of the metric comparisons involve traditional 
osteometries and associated indices. For these, the values for Gough's 
Cave 1, the total 'Mesolithic' sample, and the male Mesolithic 
samples are provided as mean ± standard deviation in the appropriate 
text position. Except for Gough's Cave 1 , right and left values were 
averaged prior to computing the sample summary statistics. 

In addition, it is appropriate to include cross-sectional geometric 
parameters (cross-sectional areas and second moments of area) into 
the description and analysis of the long bone diaphyses of fossil 
hominids. Consequently, these data are included for Gough's Cave 1 
in the description of the femoral and tibial diaphyses (Tables 6, 8, 12, 
14). Comparative data are less abundant. They have been generated 
for the full femoral and tibial diaphyses (five sections each) by S.E. 
Churchill and myself for most of the Mesolithic remains from 
Gramat, Hoedic, Rochereil and Teviec; additional data for the proxi- 
mal and midshaft femur and midshaft tibia are available from B. Holt 
(1999) (see Tables 7, 9, 13, 15 for sample sizes). 

All of the Gough's Cave 1 and most of the comparative Mesolithic 
cross sections were reconstructed using transcriptions of the subpe- 
riosteal contours and interpolations of the endosteal contours from 
anterior, posterior, medial and lateral cortical thicknesses. These 
were done at 20%, 35%, 50%, 65% and 80% of biomechanical 
length, as preservation permitted. The subperiosteal contours were 
taken using silicone putty molds [using Cuttersil Putty Plus (Heraeus 
Kulzer Inc.)] perpendicular to the diaphyseal axis, which were then 
transcribed onto paper. Cortical thicknesses were measured on antero- 
posterior and medio-lateral radiographs of the diaphyses, correcting 
for parallax using the subperiosteal diameters. The endosteal con- 
tours were manually interpolated using the cortical thickness 
rectangle to limit their extent and the subperiosteal contours as a 
guide. The resultant cross sections were digitized and cross-sec- 
tional geometric parameters were computed using a PC-DOS version 



© The Natural History Museum. 2003 



E. TRINKAUS 



(Eschman, 1992) of SLICE (Nagurka & Hayes, 1970). All sections 
were digitized twice and the results averaged. 

From this, six primary measurements were computed. These 
included total subperiosteal (TA) and cortical (CA) areas, from 
which medullary area (MA) can be computed, as well as the second 
moments of area relative to the antero-posterior (I ) and medio- 
lateral (I ) axes, the maximum second moment of area (I ), and the 
perpendicular to I (I ). The polar moment of area (J, or I ), a 
measure of torsional rigidity and overall strength, is the sum of any 
two perpendicular second moments of area (usually I + I , but 
also equal to I x + I ) 

For a few of the Mesolithic comparative specimens, subperio- 
steal contour molds were unavailable. For these, the 
cross-sectional parameters were computed using standard ellipse 
formulae (Runestad et al., 1993) from the subperiosteal diameters 
and cortical thicknesses. Given the antero-posterior and medio- 
lateral orientations of the radiographs, the resultant cross-sectional 
measures include only cross-sectional areas and antero-posterior 
(I ) and medio-lateral (I ) second moments of area, plus the polar 
moment of area computed as the sum of I x and I . For these, the 
resultant computed values were corrected for parallax and non- 
ellipse shapes of the cross-sections using least squares regressions 
between the radiographically determined measurements and the 
cross-sectional values obtained from digitizing the same sections 
of the other Mesolithic femora or tibiae. 

To assess proportions in the Gough*s Cave 1 diaphyses using 
cross-sectional parameters, three shape indices were computed, 
percent cortical area (%CA: (CA/TA) x 100), I/I (as a ratio) and 
I /I . (also as a ratio). The last two assess diaphyseal shape at the 
cross section locations, the former with respect to the anatomical 



axes and the latter with respect to the axis of maximum bending 
rigidity. The second is especially appropriate in the proximal femo- 
ral diaphysis and along the tibial diaphysis, given varying degrees of 
torsion in the proximal epiphyses of these bones. 

To assess robusticity, and hence to scale the cross sectional 
parameters to appropriate body size and beam characteristics (Ruff 
etal., 1993), cortical areas (as a reflection of axial loading levels) and 
polar moments of area (as a measure of resistance to bending and 
torsional loads) should be plotted against appropriate powers of long 
bone lengths adjusted for variance in body laterality and crural 
indices. Cortical areas should scale to body mass, which is propor- 
tional to femoral length cubed (Ruffe? a/.. 1993). Polar moments of 
area should scale to body mass times beam length, all raised to the 
four-thirds power (Ruffe? al., 1993). In other words, for the femur J 
°c (FL 3 xFL) 4/, = FL 16/3 andforthe tibia J°c (FL 5 xTL) 4/, = FL 4 xTL 4 '\ 

However, given the apparently similar degrees of body laterality 
and crural indices across these European terminal Upper Paleolithic 
and Mesolithic samples, as is expected by theoretical considerations 
(Ruff, 1991)and supported by current data(Holliday, 1995;Holliday 
& Churchill. 2003). it is appropriate to simply scale logged cortical 
areas and logged polar moments of area against logged bone length. 
Since this approach avoids determining the actual allometric scaling 
coefficient for each of these bones, it is employed here. 

In addition, even though comparative data are not available, 
metatarsal midshaft cross-sectional geometric measures are pro- 
vided (Table 20). They were computed from radiographically 
determined subperiosteal diameters and cortical thicknesses using 
ellipse formulae (Runestad et al., 1993) after the radiographic meas- 
urements were corrected for parallax using the osteometrically 
determined diaphyseal diameters. 




Fig. 1 Ventral (left) and dorsal (right) views of the sacrum; x 0.75. 



GOUGH'S CAVE 1: STUDY OF PELVIS AND LOWER LIMBS 



PELVIC REMAINS 



Inventory 

The pelvis is conserved fully articulated, with the two coxal bones in 
articulation with the sacrum and with each other at the pubic sym- 
physis (Figs 1-5). As a result, overall dimensions and proportions are 
readily ascertainable, but the configurations of the sacroiliac and 
pubic symphyseal surfaces are not observable. In addition, there is a 
bolt transversely through the sacroiliac articulations, the S2 and the 
dorsal ilia which maintains the pelvis in articulation. It is only 
apparent on the external ilia just dorso-cranial of the dorsal greater 
sciatic notches. 

Despite minor abrasion to several of the margins, there is no 
apparent distortion to any of these bones, and adhering matrix is thin 
and scattered. This makes morphological observations on them 
highly reliable. 

Sacrum. The sacrum is largely complete from the cranial S 1 to the 
caudal S5. with minor abrasion to several of the edges. The primary 
areas of abrasion are across the sacral promontory producing a 
rounded margin, and on most of the SI cranial disk surface and the 
cranial surfaces of the alae. There is also minor surface bone loss 
along the edges of the sacro-iliac articulations, but it is largely 
obscured by their articulations with the ilia. There is also a rounded 
hole dorso-ventrally through the S2 body, the result of a bolt placed 
through it for the previous mounting of the articulated skeleton in the 
Gough's Cave Museum. 

Right Coxal Bone (No. 1 . 1/23). The right coxal bone is essentially 
intact. There is abrasion to the ventro-caudal ischio-pubic ramus 
margin just ventral of the ischial tuberosity, to the internal margin of 
the mid iliac crest, and along the superior auricular margin extending 
on to the dorsal arcuate line. In addition, the middle of the iliac fossa 
has an area of adhering matrix and a small hole (maximum diameter: 
6.5mm) in the middle of that area. All of the iliac crest is present, 
even though it is partially fused. 

Left Coxal Bone (No. 1.1/24). The left coxal bone is similarly 
intact without distortion. It shares the same abrasion to the ventro- 
caudal margin of the ischio-pubic ramus just ventral of the ischial 
tuberosity and to the cranial margin of the auricular surface and 
adjacent arcuate line. In addition, there is a notch of bone missing 
from the ventral ilium just below the anterior superior iliac spine, and 
there is a large hole (31.8mm dorso- ventral and 23.0mm cranio- 
caudal) in the middle of the iliac fossa. The iliac crest is present 
ventrally, but it was (at least partially) unfused between the iliac 
pillar and the iliac tuberosity and is absent from that portion of the 
ilium. 

Pelvic Morphology 

Sacrum (Table 1 ; Fig. 1 ). The Gough's Cave 1 sacrum retains five 
clear sacral vertebrae. In this they follow the pattern of the majority 
of recent humans (Schultz, 1930). Despite damage in the regions of 
the auricular surfaces, it appears that the lateral portions of the 
sacrum and their dorsal neural arches were fully fused at the time of 
death. However, the bodies remain largely separate across their 
ventral margins. The degree of fusion of the sacral bodies and the 
pattern of fusion (from caudal to cranial) primarily reflects the young 
adult age of the individual and not an unusual pattern or degree of 
sacral fusion. 

The ventral length of the Gough's Cave 1 sacrum of ca. 123.7mm 
is large for a recent human (Radlauer, 1908). In combination with a 
mean femoral bicondylar length of 436.0mm, it provides a length 



Table 1 Osteometries of the Gough's Cave 1 sacrum. 

Ventral height chord (M-2) 1 

Ventral height arc (M-l) 

Ventral SI height chord 2 

Ventral S2 height chord 

Ventral S3 height chord 

Ventral S4 height chord 

Ventral S5 height chord 

Dorsal height chord (M-3) 

Antero-cranial breadth (M-5) 

Mid sacral breadth (M-9) 

Base dorso-ventral diameter (M-18) 

Base transverse diameter (M-l 9) 

Base sagittal angle' 

Base/S 1 sagittal angle 4 

Canal dorso-ventral diameter (M-16) 

Canal transverse diameter (M-17) 



(123.7) 

(131.5) 

(32.2) 

30.2 

25.9 

21.8 

19.5 

124.7 

(110.0) 

83.0 

(29.0) 

46.2 

81° 

60° 

17.6 

30.6 



1 (M-xx) refers to the equivalent measurement in R. Martin's Lehrbuch der 
Anthropologie (see Brauer, 1988). 

2 Cranio-caudal distance between the cranial and caudal margins of each ventral 
body. 

3 The angle, in the median sagittal plane, between the tangent to the SI vertebral disk 
surface and the ventral height chord from SI to S5. 

4 The angle, in the median sagittal plane, between the tangent to the SI vertebral disk 
surface and the ventral surface of S 1 . 

index of ca.28.4. This value high for a recent human sample (Warren, 
1897; Trinkaus, 1983) but it is only slightly above a Mesolithic 
sample mean (27.6 ± 2.2, N = 14) and very close to the mean of a 
Mesolithic male sample (28.0 ± 2.2, N = 1 1 ). 

The maximum antero-cranial breadth of the Gough's Cave 1 
sacrum (ca. 11 0.0mm) is moderate compared to other Mesolithic 
remains, and it provides an index against ventral height of 88.9. This 
value is only slightly below that of a highly variable Mesolithic 
sample (91.4 ± 8.9, N= 16) and removing the three females from the 
sample moves the mean close to the Gough's Cave 1 value (Mesolithic 
males: 89.8 ± 8.4, N = 13). 

The sacrum presents a modest degree of ventral concavity, as is 
indicated by an index of the ventral chord to the ventral arc of 
ca.94. 1 . This index is well above the mean of a Euroamerican male 
sample [85.8 ± 4.7, N = 50 (Tague, 1989)]. However, it is quite close 
to means of 93.3 for both Mesolithic samples (pooled sex sample: ± 
2.5, N = 9; males: ± 2.6, N = 8). Most of the curvature present is in 
the vicinity of S4, with only a slight concavity cranial of the S3/S4 
articulation. 

The sacral foramina are all present and prominent. They are 
slightly larger on the left side, primarily in cranio-caudal height, but 
present no unusual features. 

The cranial surface of the SI is notable for the degree of caudal 
slope of the alae, from the lateral margins of the S 1 body to the 
cranial margins of the auricular surfaces (or their estimated positions 
given damage). The degree of downward slope is indicated by a 
cranio-caudal distance of 2 1 .0mm between the promontory and a 
line between the intersections of the arcuate lines and the auricular 
surfaces. In a parallel way, the S5 body extends caudally from its 
lateral portions, down to a clearly delimited body surface for the Cx 1 
articulation. 

The sacral hiatus extends cranially to the level of the S3/S4 
intervertebral body articulation. In two recent human samples, 
Euroamericans and Afroamericans, about a third of the individuals 
have the hiatus extend cranially to the cranial S4 or above [34.3%, N 
= 519 and 30.4%, N = 694 respectively (Trotter & Lanier, 1945)], 
making this pattern in Gough's Cave 1 relatively common. 

Ilia (Table 2; Fig. 2). The Gough's Cave 1 ilia present relatively 
smooth surfaces but with generally clear markings for the various 



E. TRINKAUS 




Fig. 2 Dorso-lateral views of the left and right ilia and ischia, with the caudal sacrum; x 0.44. 



Table 2 Osteometries of the Gough's Cave 
sciatic notches. 



ilia, acetabulae and greater 





Right; 1.1/23 


Left; 1.1/24 


Iliac blade height (M-10) 


100.0 


100.9 


Iliac blade depth (M-ll) 


9.2 


(8.0) 


Superior iliac breadth (M-12) 


166.0 


164.5 


Inferior iliac breadth 1 


119.4 


(113.0) 


Arcuate line chord 2 


- 


59.0 


Arcuate line subtense 3 


- 


3.5 


Acetabular height 4 


53.8 


53.5 


Acetabular depth 5 


24.0 


26.1 


Acetabulo-sciatic breadth'' 


35.6 


35.2 


Greater sciatic notch height 7 


54.6 


(50.0) 


Greater sciatic notch breadth 8 


40.9 


42.2 



1 Maximum direct length around the anterior inferior iliac spine and the posterior 

inferior iliac spine. 

: Anterior margin of the auricular surface to the point on the arcuate line where a 

line, perpendicular to the arcuate line and passing through the depth of the psoas 

groove below the anterior inferior iliac spine, meets the arcuate line (Ruff, 1995). 

' Maximum subtense from the arcuate line chord to the arcuate line. 

1 Acetabular margin height from the margin adjacent to the anterior inferior iliac 

spine to the most distant point on the inferior acetabulum, measuring only on the 

subchondral bone of the acetabulum proper. 

5 Maximum depth from the height chord to the subchondral bone. 

"Miminum distance from the postero-lateral margin of the acetabulum to the ischial 

margin of the greater sciatic notch. 

7 Direct distance from the middle of the ischial spine to the middle of the posterior 

inferior iliac spine. 

* Direct distance from the middle of the posterior inferior iliac spine to the posterior 

ischial margin of the greater sciatic notch, taken perpendicular to the ischial margin 

between the notch itself and the ischial spine. 



muscular attachments. Externally, the gluteal abductor surfaces 
show little relief. One can perceive a M. gluteus minimus line curving 
from the iliac crest to the greater sciatic notch region, and there is 
smooth vertical ridging on the surface dorsal of that line. Internally, 
there is erosion on both sides but the preserved areas are evenly 
concave and smooth. The iliac portions of the arcuate lines are 
rounded angles from the acetabular area to the auricular surfaces. 

The cranial surfaces of the greater sciatic notches are smooth 
bilaterally. The right one, however, presents a prominent pre-auricu- 
lar sulcus, with some rugosity but mostly resorptive bone. The 
well-preserved left one, in contrast, is smooth with no trace of a pre- 
auricular sulcus. 

The iliac crest is moderately developed where it is preserved, with 
minimal rugosity. Similarly, the anterior superior iliac spine is 
modest in its development, producing only a small concavity in 
lateral view between it and the anterior inferior iliac spine (at least on 
the right side, where the bone is intact). The anterior inferior iliac 
spines are prominent and thick, but there is no lateral rotation of the 
spines or internal concavity to them. Yet, they are accompanied by a 
distinct sulcus between them and the acetabular margin, ca. 1.5mm 
wide on each side. The attachment area for the long head of M. rectus 
femoris is evident but not accompanied by marked rugosity or 
surface bone resorption. 

Ischia (Table 3; Figs 2, 3). The ischial tuberosities are generally 
smooth with prominent proximal depressions for the insertions of 
Mm. semimembranosus . They are clearly differentiated from their 
adjacent acetabular margins as well as from the ventro-lateral surface 



GOUGH'S CAVE 1: STUDY OF PELVIS AND LOWER LIMBS 




Fig. 3 Ventro-lateral view of the right ischiopubic region, with ventro- 
medial view of the left ischiopubic ramus; x 0.6. 

Table 3 Osteometries of the Gough's Cave 1 ischio-pubic regions. 



Pubic length 1 


90.4 


90.2 


Acetabulo-symphyseal length 2 


65.5 


70.1 


Ventral pubic ramus thickness 3 


11.0 


11.4 


Symphyseal height (M-18) 


41.7 


42.4 


Symphyseal breadth (M-19) 


20.2 


21.7 


Symphyseal body breadth 4 


23.7 


24.0 


Obturator foramen length (M-20) 


62.2 


61.5 


Obturator foramen breadth (M-21) 


36.6 


37.2 


Ischial length 5 


87.0 


86.8 


Ischial tuberosity breadth 6 


27.2 


28.5 


Ischio-pubic ramus height 7 


- 


16.2 


Ischio-pubic ramus thickness 8 


- 


8.2 


Ischio-pubic chord'' 


102.8 


(102.0) 


Arcuate line chord 10 


- 


125.0 



1 Direct distance from the middle of the acetabulum to the medial symphysis. 

2 Direct distance from the medial symphysis to the nearest point on the acetabular 
marrgin (McCown & Keith, 1939). 

3 Minimum thickness from the sulcus for the obturator vessels and nerve to the 
middle of the cranial surface of the superior pubic ramus (Trinkaus, 1983). 

4 Minimum distance from the middle of the pubic symphysis to the adjacent 
obturator foramen margin. 

5 Mid-acetabular point on the superior margin of the acetabular notch to the furthest 
point on the ischial tuberosity. 

* Maximum breadth of the muscle attachment area on the tuberosity. 

' Minimum dimension of the ramus, measured in an supero-dorsal to infero- ventral 

direction, parallel to the ventro-lateral surface of the ramus. 

8 Minimum dimension of the ramus, measured perpendicular to the ramus height. 

'Direct distance between the dorsal end of the ischial spine and the inferior margin 

of the pubic symphysis [Tague (1989) measurement MD], 

10 Direct distance along the arcuate line from where it meets the anterior margin of 

the auricular surface to where it meets the pubic symphysis [Tague (1989) 

measurement KO]. 

of the ischium along the obturator foramen. There is a slight develop- 
ment of ridged bone between the dorso-cranial comers of the 
tuberosities and the ischial spines for the bursae of each Mm. 
obturator internus, especially on the left side. However, the sulci for 
each Mm. obturator internus do not impinge on the tuberosities, as in 
many recent and Late Pleistocene humans (Trinkaus, 1996). This is 
accompanied by a strong lateral rotation of the tuberosities, such that 
their primary muscular surfaces are almost in the same planes as the 
external ilia. 

The tips of the ischial spines are absent, but they appear to have 
been curved inwards and moderately robust. 



Pubic bones (Table 3; Fig. 3). The pubic bones present prominent 
pubic tubercles for the inguinal ligaments, accompanied by clear, 
angled but not cresting pectineal lines, extending from adjacent to 
the acetabulae to the symphysis. The ventral margins of the superior 
pubic rami are moderately thick (11.0 and 11.4 mm), and end 
ventrally in rounded but downwardly curved margins. The symphy- 
seal bodies are narrow. 

The internal surfaces of the symphyseal bodies and the ischio- 
pubic rami are smooth with only a hint of musculo-ligamentous 
attachments, but the cranial two-thirds of the external ischio-pubic 
rami have strong muscular markings and are ventrally flared. 

Acetabulae (Table 2). There is little of note on the acetabulae 
except for a large pit on each of the subchondral bone surfaces in the 
middle of the weight bearing portion (the middle of the iliac portion 
between the anterior inferior iliac spine and the iliac pillar). The 
details of it are obscured on the left side by adhering matrix, but on 
the right side it is accompanied by a large vascular groove between 
it and the acetabular notch plus a smaller pit 19.0mm ventral of it 
immediately below the anterior inferior iliac spine. 

Pelvis as a Whole (Table 4; Figs 4, 5). The articulated Gough's 
Cave 1 pelvis presents a largely symmetrical outline. The only real 
right-to-left contrast is in the degree of iliac flare, in which the left 
ilium is more laterally and less vertically oriented. The only other 
visual difference, the apparently more open sub-pubic angle on the 
right side, is the product of postmortem abrasion to the right ischio- 
pubic ramus. 

The completeness of the Gough's Cave 1 pelvis permits compari- 
sonsof some 'obstetric' dimensions to those of at least recent human 
samples (Tague, 1989). In particular, comparisons are made to 
Euroamerican males, matching sex and approximate geographic 
origin. The pelvic funneling index of Gough's Cave 1 (outlet (bi- 
tuberous) breadth vs. inlet breadth: 79.2) is essentially the same as 
the mean of the recent Euroamerican male sample (78.8 ± 7.9, N = 
50), and similar to the means of Afroamerican and Amerindian male 
samples and well below the means of similar female samples (Tague, 
1989). However, the inlet, midplane and outlet shape indices (dorso- 
ventral vs. transverse diameter) of Gough's Cave 1 (100.0, 115.0and 
104.2 respectively) contrast with those of the Euroamerican male 
sample (79.0 ± 7.9, 133.4 ± 6.9, 111.1 ± 14.1; N = 50). In this, 
Gough's Cave 1 has a much rounder pelvic inlet, one which is 
hyperfemale. Its midplane index is low for either males or females, 
and its outlet proportions are between the means of the Euroamerican 
male and female samples. These proportions therefore combine with 
several other aspects of its pelvic morphology in indicating a rela- 
tively female-like but male pelvis. 

Table 4 Osteometries of the Gough's Cave 1 articulated pelvis. 



Pelvic inlet antero-posterior diameter (M-23) 
Pelvic midplane antero-posterior diameter 1 
Pelvic outlet antero-posterior diameter 2 
Bi-iliac breadth (M-2) 
Pelvic inlet transverse breadth (M-24) 
Articular bi-acetabular breadth (M-7) 
Minimum bi-acetabular breadth (M-7(l)) 
Bi-spinous breadth (M-8) 
Bi-tuberous (outlet) breadth 1 
Sub-pubic angle (M-33) 



125.0 

112.0 

103.2 

274.0 

125.0 

126.0 

115.0 

97.4 

99.0 

64° 



1 Direct distance from transverse ventral line between fourth and fifth sacral vertebral 
bodies to dorsomedial margin of the inferior pubic symphysis [Tague (1989) 
measurement CD). 

2 Direct distance from the ventral apex of the fifth sacral vertebra to the dorso-medial 
margin of the inferior pubic symphysis [Tague (1989) measurement DE]. 

1 Minimum distance between the two ischial tuberosities. 



E. TRINKAUS 




Fig. 4 Cranial view of the articulated Gough's Cave 1 pelvis; x 0.6. 




Fig. 5 Ventral view of the articulated Gough's Cave 1 pelvis; x 0.6. 



GOUGH'S CAVE 1: STUDY OF PELVIS AND LOWER LIMBS 



FEMORA 



Inventory 

Right (No. 1.1/35) 

The right femur is essentially complete. There is minor damage to 
the anterior head margin over an area 15.3mm proximo-distal by 
17.0mm antero-posterior, and there was abrasion to the medial 
margin of the medial condyle which obscures the medial articular 
margin. In addition, there is matrix adhering to the intertrochanteric 
crest combined with surface bone damage. 



Left (No. 1.1/34) 

The left femur is a complete bone with trivial edge abrasion to the 
condyles, and a loss of surface bone to the postero-proximal head 
over an area 2 1 .5 by 19.0 mm. 

Morphology 

The Gough's Cave 1 femora are long, slender and relatively straight 
bones, with moderate muscular markings. This is combined with 
moderately sized articulations (Figs 6, 7). 

The maximum and bicondylar lengths of the two bones differ 




Fig. 6 Anterior (left) and posterior (right) views of the Gough's Cave I femora; x 0.4. 



E. TRINKAUS 




Fig. 7 Medial (left) and lateral (right) views of the Gough's Cave 1 femora; x 0.4. 



slightly, with the right interarticular lengths being ca.6.0mm longer 
(Table 5). However, all of this difference is contained within the 
proximal epiphyses, since the bicondylar trochanteric lengths are 
identical and the left maximum trochanteric length is slightly longer. 

Diaphyses (Tables 5, 6, 8; Figs 6, 7) 

The diaphyses are straight medio-laterally with the minimum breadth 
near midshaft. The femora have moderate anterior curvature, as is 
indicated by subtense/chord indices of 3 . 1 and 2.9 versus 3 .4 ± 0.7 (N 
= 16) for a Mesolithic sample and 3.5 ± 0.6 (N = 1 0) for a Mesolithic 
male sample. It is produced primarily by an anterior angulation in the 



mid-proximal diaphysis with relatively straight more proximal and 
distal diaphyseal profiles. As a result, the positions of the maximum 
subtenses are 43.9% and 39.7% of the chords from their proximal 
ends, values which are only slightly below the means of variable 
Mesolithic (46.0 ± 9.0, N = 15) and Mesolithic male (47.1 ± 10.4, N 
= 9) samples. 

The diaphyses exhibit clear asymmetry near midshaft with the 
right side being larger. This is reflected in larger right side midshaft 
diameters (Table 5). It is more evident in cross-sectional measures 
(Tables 6, 8), which exhibit a 6.7% asymmetry in cortical area and a 
16.9% asymmetry in the polar moment of area [% asymmetry = 



GOUGH'S CAVE 1: STUDY OF PELVIS AND LOWER LIMBS 



Table 5 Length measurements of the Gough's Cave 1 femora and 
osteometries of the femoral diaphyses . 



Right 



Left 



Maximum length (M-l) 

Bicondylar length (M-2) 

Trochanteric length (M-3) 

Bicondylar trochanteric length (M-4) 

Biomechanical length 1 

Midshaft antero-posterior diameter (M-6) 

Midshaft medio-lateral diameter (M-7) 

Midshaft circumference (M-8) 

Subtrochanteric antero-posterior diameter (M- 10) 2 

Subtrochanteric medio-lateral diameter (M-9) 

Subtrochanteric circumference 

Anterior curvature chord (M-27) 

Anterior curvature subtense 

Anterior curvature subtense position 3 



1 Distance parallel (o the diaphyseal axis between the intersection of that axis with 
the proximal neck (just medial of the greater trochanter) and the average of the 
positions along the diaphyseal axis of the distal condyles (Ruff & Hayes, 1983). 

2 The subtrochanteric diameters are taken as the maximum medio-lateral dimension 
(usually close to the antero-medial to postero-lateral plane of anteversion) and the 
antero-posterior diameter perpendicular to that medio-lateral one. 

3 Distance from the proximal end of the chord to the position of the maximum 
subtense. 



Table 6 Cross-sectional second moments of area of the Gough's Cave 1 
femoral diaphyses (in mm 4 and degrees). 



443.0 


437.5 


439.0 


433.0 


423.0 


424.5 


413.0 


413.0 


415.3 


408.0 


33.7 


30.2 


24.8 


23.8 


92.0 


85.5 


28.2 


27.6 


31.5 


31.7 


94.0 


93.0 


303.0 


295.0 


9.5 


8.5 


133.0 


117.0 



Right 



Left 



20% AP second moment of area (I x ) 
20% ML second moment of area (I ) 
20% Maximum second moment of area 
20% Minimum second moment of area 
20% Polar moment of area (J) 
20% Angle of I max (theta) 
35% AP second moment of area (I x ) 
35% ML second moment of area (I ) 

y 

35% Maximum second moment of area 

35% Minimum second moment of area 

35% Polar moment of area (J) 

35% Angle of I max (theta) 

50% AP second moment of area (I x ) 

50% ML second moment of area (I ) 
y 
50% Maximum second moment of area 

50% Minimum second moment of area 

50% Polar moment of area (J) 

50% Angle of I max (theta) 

65% AP second moment of area (I ) 

65% ML second moment of area (I ) 
y 
65% Maximum second moment of area 

65% Minimum second moment of area 

65% Polar moment of area (J) 

65% Angle of 1^ (theta) 

80% AP second moment of area (I x ) 

80% ML second moment of area (I ) 
y 
80% Maximum second moment of area 

80% Minimum second moment of area 

80% Polar moment of area (J) 

80% Angle of I (theta) 



(I ) 



tfnJ 

(IJ 



(U 



(I ) 



A—) 
(I J 



35927.1 
39249.1 
40399.9 
34776.2 
75176.1 

27° 
35539.9 
20315.4 
35629.1 
20226.3 
55855.4 

94° 

39160.3 
21418.6 
40129.3 
20449.5 
60578.8 

103° 
39874.0 
23495.7 
41309.5 
22060.2 
63369.7 

106° 
30878.2 
31956.3 
33380.3 
29454.2 
62834.5 

37° 



29219.4 
35295.2 
36884.4 
27630.1 
64514.5 

25° 
28073.8 
20642.1 
28972.0 
19743.9 
48715.9 

108° 
28671.0 
21693.3 
30867.6 
19496.7 
50364.3 

116° 
29467.0 
22797.5 
31796.8 
20467.7 
52264.5 

117° 
30129.1 
31368.4 
34249.7 
27247.8 
61497.5 

40° 



((max - min)/max) x 100]. In contrast, in the proximal diaphysis, 
even though the right side continues to be larger, the level of 
asymmetry is much less, with cortical area exhibiting 4.8% asymme- 
try and the polar moment of area providing only a 2. 1 % contrast. 

The lineae asperae are smooth along the entire lengths of the bones, 
which is possibly the product of the young adult age of the individual. 
They are along prominent pilasters for most of the middle half to two- 



thirds of the diaphysis. The pilasters are formed by antero-posteriorly 
convex medial surfaces but distinctly antero-posteriorly concave 
lateral surfaces. This results in a sulcus along the lateral pilaster 
especially in the midshaft region. The lineae asperae taper off gradually 
disto-medially, ending in moderate adductor tubercles. 

The prominence of the Gough's Cave 1 pilasters is evident by their 
pilastric indices of 1 35.9 and 126.9. Both of them, and especially the 
right one, are well above the means of Mesolithic (109.5 ± 10.6, N = 
52) and Mesolithic male (1 13.0 ± 9.2, N = 34) samples. This is 
further and better illustrated, albeit with smaller comparative samples, 
by the Gough's Cave 1 I /I and I /I ratios (Table 7) along the 

J ° x y max mm v / o 

middle third of the diaphysis (the 35%, 50% and 65% sections). 
Again, the right femur is more pilastric than the left one. 

Proximally, the markings in the pectineal region are very light, and 
they are bordered laterally by small but rugose gluteal tuberosities 
(Fig. 8). Neither gluteal tuberosity is projecting or concave, and there 
is no trace of hypotrochanteric fossae. The right tuberosity fades out 
proximally, but the left one leads to a small tubercle at the proximo- 
distal level of the lesser trochanter. The modest dimensions of the 
Gough's Cave 1 gluteal tuberosities are demonstrated by compari- 
sons of their maximum breadths. The absolute breadths (8.2 and 8.4 
mm) are below the means Mesolithic (1 1.6 ± 1.9 mm, N = 17) and 
especially Mesolithic male ( 12.3 ± 1.9 mm, N = 10) samples. This is 
further illustrated by indices between the gluteal tuberosity breadths 
and the geometric means of the associated subtrochanteric diaphy- 
seal diameters - the mean of the resultant values of 27.5 and 28.4 for 
Gough's Cave 1 are 1.91 and 1.98 standard deviations below the 
means respectively of Mesolithic (42.3 ± 7.5, N = 1 7) and Mesolithic 
male (43.0 ± 7.6, N = 10) samples. 

The gluteal buttresses are pronounced, with distinct sulci formed 
anteriorly and posteriorly. The right one is covered posteriorly by the 
gluteal tuberosity, but the left gluteal tuberosity covers only the 
medial half of the buttress. Nonetheless, the subtrochanteric diaphy- 
ses of Gough's Cave 1 are relatively round compared to those of most 
Mesolithic femora. Its meric indices of 89.5 and 87.1 are 2.49 and 
2.84 standard deviations above the means respectively of pooled 
Mesolithic (74.6 ± 5.5, N = 85) and Mesolithic male (75.8 ± 4.4, N 
= 50) samples. However, the large sample Muge has significantly 
lower meric indices (73.0 ± 4.8, N - 55) than the remainder of the 
Mesolithic sample (P < 0.001). Yet even using only non-Muge 
Mesolithic remains for the comparison still places the Gough's Cave 
1 femora 1.93 standard deviations from the mean (77.5 ± 5.6, N = 
30). Similarly, the proximal diaphyseal (80%) I /I s ratios are 2.43 
and 2.70 standard deviations below the means of respective Mesolithic 
samples (Table 7). 



Table 7 Comparative femoral second moment of area diaphyseal shape 
indices, I/I and I max /I min , for Gough's Cave 1 and Mesolithic samples. 
For Mesolithic samples, mean ± SD is given. 



I/I 



Gough's Cave 1: 
Right; Left 



Mesolithic Sample Mesolithic Males 



20% 
35% 
50% 
65% 
80% 

I max /I m 

20% 
35% 
50% 
65% 



0.92; 0.83 
1.75; 1.36 
1.83;1.32 
1.70; 1.29 
0.97; 0.96 



1.16; 1.33 
1.76; 1.47 
1.96; 1.58 
1.87; 1.55 
1.13; 1.26 



0.66 ±0.08 
1.01 ±0.15 
1.18 ±0.23 
1.00 + 0.21 
0.77 ±0.20 



1.58 + 0.19 
1.20 + 0.12 
1.33 ±0.19 

1.32 ±0.26 
1.90 ±0.29 



N= 16 
N=14 
N = 55 
N= 15 

N = 52 



N= 13 

N= 11 
N = 45 
N= 12 
N = 41 



0.68 ± 0.07; N= 10 
1.02±0.20;N = 8 
1.20 ± 0.22; N = 37 
1.08 ±0.17; N = 9 
0.79 ± 0.19; N = 34 



1.54 ±0.18; N = 8 
1.26 ±0.12; N = 6 
1.33 ± 0.19; N = 30 
1.25±0.13;N = 7 
1.87 ± 0.25; N = 27 



10 



E. TRINKAUS 




Fig. 8 Proximo-posterior view of the Gough's Cave 1 femora: x 0.8 (enlargement of Fig. 6. top right). 



Table 8 Cross-sectional area measures of the Gough's Cave 1 femoral 
diaphyses (in mm 2 ). 



20% Total area (TA) 
20% Cortical area (CA) 
20% Medullary area (MA) 
35% Total area (TA) 
35% Cortical area (CA) 
35% Medullary area (MA) 
50% Total area (TA) 
50% Cortical area (CA) 
50% Medullary area (MA) 
65% Total area (TA) 
65% Cortical area (CA) 
65% Medullary area (MA) 
80% Total area (TA) 
80% Cortical area (CA) 
80% Medullary area (MA) 



Right 



806.3 
390.4 
415.9 
612.0 
390.4 
221.6 
595.3 
507.2 

88.1 
618.8 
544.1 

74.7 
668.0 
433.4 
234.6 



Left 



751.3 
343.3 
408.0 
562.8 
417.9 
144.9 
552.9 
473.3 

79.6 
572.5 
479.5 

93.0 
668.8 
412.7 
256.1 



The overall robusticity of the femoral diaphyses can be compared 
to those of relatively large samples of European Mesolithic femora 
using its midshaft external diameters [((AP x ML)" : / bicondylar 
length) x 100]. The resultant indices are 6.6 and 6.2 for Gough's 



Table 9 Comparative femoral percent cortical area (%CA = (CA/TA) x 
100) for Gough's Cave 1 and Mesolithic samples. 



Gough's Cave 1: 
Right; Left 



Mesolithic Sample Mesolithic Males 



20% 
35% 
50% 
65% 
80% 



48.4; 45.6 
63.8:74.3 
85.2; 85.6 
87.9; 83.6 
64.9;61.7 



48.2±4.1;N=16 
67.3 ± 5.8; N= 14 
79.0 ± 6.9; N = 55 
86.3 ± 2.6: N= 15 
77.6 ± 8.3: N = 52 



47.3 ± 3.6; N= 10 
65.1 ±3.3; N = 8 
77.9 ± 6.3; N = 37 
85.7±2.7;N = 9 
78.1 ± 6.8; N = 34 



Cave 1 , the right one of which is on the means of Mesolithic (6.5 ± 
0.4, N = 47 ) and Mesolithic male (6.6 ± 0.4, N = 3 1 ) samples and the 
left one only slightly below them. In this, the large Muge sample has 
a significantly (P = 0.002) lower mean (6.3 ± 0.2, N = 19), but its 
inclusion or deletion from the Mesolithic sample has little effect on 
the relative position of Gough's Cave 1. 

Using midshaft cross-sectional measures, the percent cortical 
areas largely cluster close to the Mesolithic comparative means; only 
the 80% one is relatively low, significantly so with respect to the 
Mesolithic male sample (Table 9). Plots of the midshaft cortical area 
and polar moment of area versus powers of femoral length (Fig. 9) 
largely support the pattern seen in the robusticity indices; Gough's 



GOUGH'S CAVE 1 : STUDY OF PELVIS AND LOWER LIMBS 



11 





't 




CO 


m 




Cl> 


CM 


< 


(O 


m 




o 




o 


o 


o 


CD 






o 




in 




r 


CO 


_i 


CO 




CD 




LD 



Table 10 Osteometries of the Gough's Cave 1 femoral epiphyses. 

Right 



Left 



□□ 



■ i, D 







5.90 5.95 6.00 

Ln Femur Length 



6.05 



6.10 



CD 


< 

+-» 
c 

CD 

E 
o 



jo 
o 



• 










o d ti 


■ 
a 


a 

a D 

n 

D 
D 

D 








n 


u 

n 


n r5L° • 

D~0 Q 












u 
n 


B □ „ O 


D 








D 




n 


D 
D 










D 
















D 






go 




□ 




D 


□ 






a 










D 






























' ' 


i i I i 




■ 


i | i i i . i | i 


' ' 


1 ' ' ' ' 1 



5.85 5.90 5.95 6.00 6.05 6.10 

Ln Femur Length 

Fig. 9 Plots of the Gough's Cave 1 femoral midshaft logged cross- 
sectional parameters vs. ln length (see text). Solid hexagons: Gough's 
Cave 1 right and left femora; gray squares: Mesolithic males; open 
squares: Mesolithic females. 



Cave 1 is well within the Mesolithic distributions, close to the 
middles of the comparative sample distributions in both measures. 

Proximal Epiphyses (Table 10; Fig. 10) 

The proximal epiphyses present overall similar morphologies but 
contrast in several aspects of their proportions, some of which are 
reflected in the slight (ca.1%) length asymmetry. The heads are 
evenly rounded and each has a large fovea capitis placed slightly 
posterior on the head. There is no trace of an Allen's fossa on either 
femur. The right head is slightly larger, especially in the cranio- 
caudal direction (Table 10), but comparisons of their head sagittal 
diameters to femoral bicondylar length produce similar indices ( 1 0.9 
& 10.7), in part due to the differences in femoral lengths. They are 
both very close to the means of Mesolithic ( 10.7 ± 0.5, N = 37) and 
Mesolithic male (10.8 ± 0.5, N = 26) samples. 

On the left femur where is it preserved, there is a large obturator 
fossa with a large pit 12.5mm deep from its posterior edge. The 
intertrochanteric crest has clearly marked fibrous spicules running 



Head-neck length (M-14) 

Anatomical biomechanical neck length 1 

Trochanteric biomechanical neck length 2 

Head sagittal diameter (M-19) 

Head vertical diameter ( M- 1 8 ) 

Neck circumference (M-17) 

Neck-shaft angle (M-29)' 

Anteversion angle (M-28) 

Greater trochanter depth (M-26( 1 )) 

Gluteal tuberosity breadth 4 

Distal epicondylar breadth (M-2 1 ) 

Bicondylar breadth 5 

Medial condylar breadth (M-2 1 c ) 

Lateral condylar breadth (M-21e) 

Bicondylar angle (M-30) 

Medial patellar projection (M-24b) 

Lateral patellar projection (M-22) 

Median patellar projection 6 

Patellar surface circumference 7 

Patellar surface breadth (M-26(3b)) 

Patellar surface depth 8 

Patellar surface depth position'' 



80.1 


79.9 


37.0 


40.0 


63.0 


67.0 


47.7 


46.3 


48.2 


46.3 


97.0 


91.5 


132° 


135° 


11° 


30° 


- 


37.3 


8.2 


8.4 


- 


78.5 


(75.6) 


76.6 


(29.4) 


29.6 


28.3 


28.6 


10° 


10° 


60.7 


58.9 


63.6 


61.9 


59.0 


58.9 


42.0 


45.0 


37.7 


39.1 


6.0 


6.5 


13.6 


13.6 



1 Distance perpendicular to the diaphyseal axis from that axis to the proximal tangent 

to the femoral head. 

- Distance perpendicular to the diaphyseal axis from the proximo-distal tangent to 

the lateral greater trochanter to the proximal tangent to the femoral head (Lovejoy et 

al, 1973). 

5 Taken in the anteversion plane of the femoral head and neck. 

4 Maximum breadth of the rugose area for the insertion of M. gluteus maximus on the 
proximal diaphysis (Trinkaus, 1976). 

5 Maximum breadth across the external medial and lateral condylar surfaces. 

6 Dorsal condylar plane to the tangent parallel to that dorsal plane within the deepest 
portion of the patellar sulcus. 

7 Articular arc in the patellar sulcus from the intercondylar margin to the proximal 
patellar surface. 

8 Maximum depth subtense to the sulcus floor, taken from the surface breadth. 

9 Position of the depth subtense from the lateral margin of the patellar surface. 



along it, from the capsular attachment, but the intertrochanteric lines 
on the anterior surfaces are faint. Both lesser trochanters are strongly 
medially projecting, and the greater trochanters have a clear lateral 
swelling below the M. gluteus medius insertion. The left greater 
trochanter also exhibits a strong beak at its proximo-medio-posterior 
margin. 

The neck shaft angles of the two femora show a moderate degree 
of asymmetry, with the left one being 3° higher. Both of these angles 
( 1 32° & 1 35°) are relatively high for foraging populations (Trinkaus, 
1993) but not exceptional for a European Mesolithic sample (125.5° 
± 5.9°, N = 22) or a male one (124.4° ± 5.4°, N = 15). However, the 
Gough's Cave 1 values nonetheless have z-scores of 1.36 and 1.68 
respectively. 

Of greater asymmetry are the anteversion angles, a modest 1 1 ° on 
the right side but a pronounced 30° on the left. This asymmetry in 
anteversion angles is reflected in direction but not in degree by the 
values for theta at the subtrochanteric level (37° and 40° respec- 
tively). In comparison, a variable Mesolithic sample provides 
anteversion angle values of 1 8.2° ± 9.0° (N = 1 8) and 1 8.3° ± 8.7° (N 
= 12) for males alone. 

The relatively high neck-shaft angles of Gough's Cave 1 contrib- 
ute in part to its relatively short biomechanical neck lengths, which 
provide indices relative to bicondylar length 8.4 and 9.2 for the 
anatomical biomechanical neck length and 14.4 and 15.5 for the 
trochanteric one. These are relative respectively to 9.6 ± 0.9 (N = 1 1 ) 
and 16.7 ± 1.0 (N= 10) for the Mesolithic sample and 9.5 ± 0.8 (N = 
7) and 16.9 ± 0.7 (N = 6) for the male sample. 



12 



E. TRINKAUS 




Fig. 10 Anterior view of the Gough's Cave 1 proximal femora; x 0.8 (enlargement of Fig. 6, top left). 



Distal Epiphyses (Table 10) 

There is little of note on the Gough's Cave 1 distal femoral epiphysis. 
The condyles are rounded with no evidence of femoral squatting 
facets. There is only a small rounded ridge along the supracondylar 
margin of the medial condyles, up to 3.5mm from the condylar 
surface on the left side where it is best preserved. 

Their bicondylar angles of 10° each are well within Mesolithic 
ranges of variation (9.7° ± 3.1, N= 19; 8.8° ± 2.5°, N = 15 for males 
only), all of which are similar to those of recent humans (Tardieu & 
Trinkaus, 1994). The one measure of distal epiphyseal size for which 
reasonable Mesolithic comparative samples exist, epicondylar 
breadth versus bicondylar length, provides an index of 18.1. It 
suggests a modest but not unusually small epiphysis, given means of 
18.8 for total (± 1.1, N = 19) and 18.9 for male (± 1.1, N = 14) 
Mesolithic samples. 



TIBIAE 

Inventory 

Right (No. 1.1/27) 

A complete bone with minor abrasion to the medial half of the 
posterior distal epiphysis and thin adhering matrix around the proxi- 
mal epiphysis. 

Left (No. 1.1/26) 

The bone retains two pieces, a proximal posterior piece of the 
diaphysis with the abraded soleal line (maximum preserved length: 
ca.80.0mm) and a diaphyseal section with all of the surfaces just 
proximal of the distal epiphysis (maximum preserved length: 
ca.50.0mm). Both of the pieces are set in a plaster and papier-mache 
reconstruction of the bone and attached to the left fibula (Fig. 1 1 ). 



Morphology 

Even though portions of both tibiae are present, meaningful morpho- 
logical observations can be made primarily on the essentially 
complete right bone. Therefore, the following comments apply to the 
right bone except where noted otherwise. 

Diaphysis (Tables 11, 12, 14; Figs 11, 12) 

The right tibial diaphysis has a gently 'S'-curved, sharply angled 
anterior crest, which is bordered medially by a smooth and slightly 
convex medial diaphyseal surface. The lateral surface is convex 
between the interosseus crest and the anterior margin and concave 
between the crest and the postero-lateral corner along the proximal 
two-thirds of the diaphysis. Distal of there, it becomes fully convex. 
The postero-medial margin is rounded proximal of the meeting of 
the soleal line with that margin, near midshaft. 

Both tibiae have a distinct but small flexor line, a crest between the 
origins of M. tibialis posterior and. M. flexor digitorum longus on the 
proximal posterior diaphysis extending distally from the soleal line. 
On the right bone, it is raised from the subperiosteal surface along 
ca.78mm distally from the soleal line. The soleal line itself is a low 
rugose area proximal of the flexor line, 4.7mm wide near the flexor 
line and 5.3mm wide more proximally. Disto-medial of the flexor 
line, the soleal line becomes a slightly raised crest, extending to the 
medial side distally. The interosseus line is a distinct crest from the 
very proximal shaft to about midshaft, at which level it becomes 
blunt and blends in with the lateral diaphysis. 

The nutrient foramen is located just medial of the flexor line, 
37.5mm distal of the intersection of the two muscular lines. 

The Gough's Cave 1 tibial diaphysis, like those of other Mesolithic 
humans, is distinctly platycnemic. Its cnemic index of 55.9 is even 
slightly below the means of Mesolithic (62.4 ± 5.8, N = 73) and 
Mesolithic male (61.8 ±6.2, N = 46) samples. Moreover, its 35% to 
80% I /I ratios are all higher than the means of Mesolithic 



GOUGH'S CAVE 1 : STUDY OF PELVIS AND LOWER LIMBS 



13 



Table 11 Osteometries of the Gough's Cave 1 right tibia. 

Maximum length (M- 1 ) 

Medial total length (M-lb) 

Medial articular length (M-2) 

Biomechanical length 1 

Midshaft antero-posterior diameter (M-8) 

Midshaft medio-lateral diameter (M-9) 

Midshaft circumference (M-10) 

Proximal antero-posterior diameter (M-8a) 

Proximal medio-lateral diameter (M-9a) 

Proximal circumference (M-lOa) 

Distal minimum circumference (M-lOb) 

Proximal epiphyseal maximum breadth (M-3) 

Medial condyle breadth (M-3a) 

Lateral condyle breadth (M-3b) 

Medial condyle depth (M-4a) 

Lateral condyle depth (M-4b) 

Tuberosity projection 2 

Medial retroversion angle (M-12) 

Lateral retroversion angle 

Medial inclination angle (M-13) 

Lateral inclination angle 

Torsion angle (M-14) 

Distal maximum breadth (M-6) 

Distal maximum depth (M-7) 

Talar trochlear articular breadth 3 

Medial talar articular depth 4 

Lateral talar articular depth 5 



385.0 
375.0 
360.0 
360.5 
34.3 
21.3 
87.5 
38.1 
21.3 
94.0 
77.0 
76.1 
34.5 
34.4 
46.7 
40.9 
48.0 
16° 
13° 
11° 
8° 
20° 
49.1 
40.3 
30.0 
26.2 
33.7 



Table 13 Comparative tibial second moment of area diaphyseal shape 
indices, I m „/I min , for Gough's Cave 1 and Mesolithic samples. 



1 The average distance, measured parallel to the diaphyseal axis, from the middle of 
the talar trochlear surface to the middle of each condyle (Ruff & Hayes, 1983). 
; The distance perpendicular to the diaphyseal axis from the antero-posterior middle 
of the condyles to the coronal plane tangent to the anterior surface of the tibial 
tuberosity (Trinkaus, 1983). 

3 The medio-lateral breadth of the talar trochlear facet from the antero-posterior 
middle of the lateral margin to the middle of the curve between the trochlear and the 
medial malleolar facets (Trinkaus, 1983; Ruff, 1990). 

4 The minimum antero-posterior dimension of the talar trochlear facet taken adjacent 
to the medial malleolus (Trinkaus. 1983; Ruff, 1990). 

s The maximum antero-posterior dimension of the talar trochlear facet taken adjacent 
to the fibular articulation (Trinkaus, 1983; Ruff, 1990). 

Table 12 Second moments of area of the Gough's Cave 1 right tibial 
diaphysis (in mm 4 and degrees). 



20% AP second moment of area (I x ) 
20% ML second moment of area (I ) 

y 

20% Maximum second moment of area (I m 

20% Minimum second moment of area (I 

20% Polar moment of area (J) 

20% Angle of I max (theta) 

35% AP second moment of area (I ) 

35% ML second moment of area (I ) 
y 
35% Maximum second moment of area (I m 

35% Minimum second moment of area (I . 

35% Polar moment of area (J) 

35% Angle of I mM (theta) 

50% AP second moment of area (1^) 

50% ML second moment of area (I ) 

50% Maximum second moment of area (I m 

50% Minimum second moment of area (I 

50% Polar moment of area (J) 

50% Angle of I nm (theta) 

65% AP second moment of area (IJ 

65% ML second moment of area (I ) 

y 

65% Maximum second moment of area (I 
65% Minimum second moment of area (I 
65% Polar moment of area (J) 
65% Angle of I max (theta) 
80% AP second moment of area (I x ) 
80% ML second moment of area (I ) 
80% Maximum second moment of area (I m 
80% Minimum second moment of area (I 
80% Polar moment of area (J) 
80% Angle of I (theta) 



14154.4 
12798.3 
15461.6 
11491.1 
26952.7 

125° 
17618.5 
12738.0 
21220.1 
9136.4 
30356.5 

119° 
29204.5 
15067.5 
33212.7 
11059.3 
44272.0 

113° 
42160.0 
16796.3 
46262.6 
12693.7 
58956.3 

111° 
58126.5 
32073.3 
65287.7 
24912.2 
90199.9 

115° 



Gough's Cave 1 



Mesolithic Sample 



Mesolithic Males 



20% 
35% 
50% 
65% 
80% 



1.35 

2.32 
3.00 
3.64 
2.62 



1.38 ± 0.25; N= 12 
2.23 ± 0.40; N = 13 
2.59 ± 0.46; N = 47 
2.89 + 0.44; N= 16 
2.44 ± 0.46; N = 1 1 



1.32 ±0.22 
2.19 ±0.34 
2.63 ± 0.47 
2.86 ±0.41 



N = 7 
N = 8 
N = 34 
N= 10 



2.47±0.53;N = 8 



Table 14 Cross-sectional areas of the Gough's Cave 
diaphysis (in mm 2 ). 

20% Total area (TA) 
20% Cortical area (CA) 
20% Medullary area (MA) 
35% Total area (TA) 
35% Cortical area (CA) 
35% Medullary area (MA) 
50% Total area (TA) 
50% Cortical area (CA) 
50% Medullary area (MA) 
65% Total area (TA) 
65% Cortical area (CA) 
65% Medullary area (MA) 
80% Total area (TA) 
80% Cortical area (CA) 
80% Medullary area (MA) 



right tibial 



480.2 
224.1 
256.1 
432.5 
299.5 
133.0 
491.1 
375.8 
115.3 
566.8 
404.4 
162.4 
793.7 
432.5 
361.2 



Table 15 Comparative tibial percent cortical area (%CA = (CA/TA) x 
100) for Gough's Cave 1 and Mesolithic samples. 





Gough's Cave 1 


Mesolithic Sample 


Mesolithic Males 


20% 


46.7 


57.3 ± 4.8; N= 15 


55.8 ± 3.9; N = 9 


35% 


69.2 


81.0 ± 3.0; N= 16 


80.8 ± 2.5; N= 10 


50% 


76.5 


84.4 ± 5.0; N = 53 


85.0 ± 4.4; N = 38 


65% 


71.4 


68.6 ± 4.5; N= 16 


68.2 ± 4.3; N= 10 


80% 


54.5 


51.2±5.0;N= 15 


51.3 ±5.4; N = 8 



samples, with the 50% and especially the 65% ratios being well 
above those Mesolithic means (Table 13). 

In terms of diaphyseal robusticity, the Gough's Cave 1 tibia on 
average is similar to those of other Mesolithic specimens. Its percent 
cortical area values are below Mesolithic means for the mid and 
distal diaphysis, but above those means in the proximal diaphysis 
(Table 15). A diaphyseal robusticity index (from the geometric mean 
of the midshaft diameters versus articular length) is 7.5 for Gough's 
Cave 1, which is very close to the means of Mesolithic (7.6 ± 0.6, N 
= 24) and Mesolithic male (7.8 ± 0.6, N = 17) samples. The plots of 
midshaft cortical area and polar moment of area versus appropriate 
powers of femoral or tibial and femoral length (Fig. 13) place 
Gough's Cave 1 clearly well within the Mesolithic ranges of varia- 
tion is slightly below a number of those specimens. 

Proximal Epiphysis (Fig. 14) 

The tibial plateau presents small intercondylar spines, a distinctly 
concave medial condylar surface, and an evenly convex lateral 
condylar surface. There is a nearly horizontal fibular facet, with its 
maximum dimension of 20.2mm approximately medio-lateral and 
the minimum diameter of 14.5mm approximately antero-posterior. 
There is a clear sulcus for the M. semimembranosus tendon, but there 
is no smoothing of the bone in the sulcus for its insertion. 

The tibial plateau is strongly rotated relative to the diaphysis, with 
a torsion angle of 20°. However, this value is close to the means of 
variable Mesolithic (22.7° ± 12.4°, N = 15) and Mesolithic male 



14 



E. TRINKAUS 




Fig. 11 Anterior (left) and posterior (right) views of the Gough's Cave 1 tibiae and fibulae, with the heavily reconstructed left tibia and fibula shown only 
in anterior view; x 0.4. 



(22.4° ± 9.9, N = 12) samples. The Gough's Cave 1 tibia also has 
clear retroversion of the condyles, with a medial retroversion angle 
of 15°. However, this value is also very close to the means of 
Mesolithic ( 15.3 ± 5. 1 °, N = 1 8) and Mesolithic male (15.0° ± 4.7°, 
N = 15) samples. All of these retroversion angles are normal for non- 
industrial recent humans (Trinkaus, 1975a). 

Similarly, the overall dimensions of the tibial plateau, quantified 
by an index of maximum breadth versus articular length of 2 1 . 1 for 
Gough's Cave 1, is normal for Mesolithic samples (22.4 ± 1.8, N - 
12, and 22.1 ± 1.5, N = 11 for males only). One feature in which the 
Gough's Cave 1 proximal tibia is further from the mean of the 
comparative samples is in relative tuberosity projection, or the 
posterior displacement of the tibial condyles from the tibial tuberos- 
ity (a measure of the M. quadriceps femoris moment arm through the 
patellar ligament). The Gough's Cave 1 value of 13.3 is significantly 



above the mean of a small Mesolithic pooled-sex sample (9.4 ± 1.4, 
N = 6), and still well above the mean of a male sample (10.5 ± 2.0, N 
= 4). 

Distal Epiphysis 

The Gough's Cave 1 tibial distal epiphysis is likewise unremarkable. 
Its talar trochlear articular surface, relative to tibial length, is similar 
in size to other Mesolithic tibiae. The index formed by the geometric 
mean of its breadth with the average of its depth measurements 
versus articular length is 8.3 - this value is close to the means for 
Mesolithic (8. 1 ± 0.6, N = 23) and Mesolithic male (8. 1 ± 0.6, N = 1 8) 
samples. 

It does present a clear lateral squatting facet, 9.0mm wide and 
3.7mm proximo-distal. There is no trace of a medial squatting facet 
or other rounding of the anterior articular margin. 



GOUGH'S CAVE 1 : STUDY OF PELVIS AND LOWER LIMBS 



15 




Fig. 12 Medial (left) and lateral (right) views of the Gough's Cave 1 right tibia and fibula: x 0.4. 



FIBULAE 



Inventory 

Right (No. 1.1/28) 

The bone consists of a proximal section with the proximal epiphysis 
and the proximal half of the diaphysis, plus a distal section with the 
distal quarter of the diaphysis and the complete distal epiphysis. The 
two pieces are joined together by a plaster reconstruction of the mid- 
distal epiphysis (minimum gap: 47.8mm), and its reconstructed 
lengths (Table 16) are based on articulation with the complete right 
tibia. 

Left (No. 1.1/26) 

Most of the diaphysis lacking both epiphyses. The epiphyses are 
reconstructed in plaster and joined to the reconstructed left tibia. 
Maximum preserved length: 279.0mm. 



Morphology 

Even though both of the diaphyses are preserved, most of the 
morphological information and all of the osteometries derive from 
the separated right fibula. Nonetheless, osteometric comparisons of 
the Gough's Cave 1 fibula are limited by poor preservation and 
limited published measurements for other European Mesolithic 
fibulae. 

Diaphyses (Table 16; Figs 11, 12) 

Both fibular diaphyses are very straight, with well formed angles for 
musculo-ligamentous attachments on all of the margins. On anterior 
view, the left one has a slight 'S' curve in the distal third, producing 
a slight lateral concavity just below midshaft - the right bone's 
reconstruction in this region (Fig. 1 1 ) may therefore be too straight. 
Even though the various angles are clearly formed, none of them 
present clear rugosities. The primary evident muscular insertions 



16 



E. TRINKAUS 





«* 




CD 




CN 


CO 


CD 


CD 




i_ 




< 


,- 


"ro 


CD 


o 




o 


en 


o 


m 



o ^ 

If) ID 



□□ 



DD 



5.65 5.70 5.75 5.80 5.85 

Ln Tibia Length 



5.90 



5.95 



^~ 












CD 








B 




CD 
























< 

*- o i 








□ 


D 


o ^~ 










D 


CD 


■ 


□ 




□ D 
Q 


D 


E 




u n 


a 


D ®D# 




«= o- 










co 


D s 










o 








D D 


CL 




%n 




□ 




50% 

10.0 


D D O n 

a d 










c 












_j 












LO 


D 
, i . i , i , i , | i i i 


1 i ' 




D 


i i i i i i 



5.65 5.70 5.75 5.80 5.85 

Ln Tibia Length 



5.90 



5.95 



Fig. 13 Plots of the Gough's Cave 1 tibial midshaft logged cross- 
sectional parameters versus loggen tibial length. Solid hexagons: 
Gough's Cave 1 right and left femora; gray squares: Mesolithic males; 
open squares: Mesolithic females. 

areas are the soleal line proximally (preserved on the right bone) and 
a broad area on the posterior midshaft (evident on both fibulae). The 
soleal line is a broad, rugose area, 13.5mm long and up to 12.7mm 
wide. It forms a slight depression and has a small lip medially. The 
right midshaft exhibits a broad rugose area ca.40.0mm long along 
the posterior surface. A similar but much less rugose area is present 
on the left diaphysis. The attachments for the distal interosseus (or 
tibio-fibular) ligaments are modest, spiraling from proximo-anterior 
to disto-posterior. 

Morphometrically, the Gough's Cave 1 right fibula is similar to 
those of other Mesolithic humans. Its midshaft maximum to mini- 
mum diameter index of 133.6 is moderately below to the means of 
highly variable samples (Mesolithic: 141.6± 17.7. N = 40; Mesolithic 
males: 140.0 ± 1 8.6, N = 22). It is slightly less robust than most other 
Mesolithic fibulae, as indicated by an index between the geometric 
mean of its midshaft diameters and maximum length (Gough's Cave 
1:3.8; Mesolithic: 4.2 ± 0.6, N = 20; Mesolithic males: 4.2 ± 0.3, N 
= 13). 

It is also possible, with smaller comparative samples, to assess its 



Table 16 Osteometries of the Gough's Cave 1 right fibula. 

Maximum length (M-l) (366.0) 

Articular length (M-la) (356.0) 

Midshaft maximum diameter (M-2) 15.9 

Midshaft minimum diameter (M-3 ) 11.9 

Midshaft circumference (M-4) 45.5 

Neck maximum diameter 13.1 

Neck minimum diameter 10.5 

Neck circumference (M-4a) 37.0 

Proximal epiphyseal medio-lateral diameter 29.8 

Proximal epiphyseal antero-posterior diameter 24.4 

Proximal tibial facet medio-lateral diameter 20.9 

Proximal tibial facet antero-posterior diameter 18.2 

Proximal articular angle ' 111° 

Distal maximum depth 2 24.9 

Distal articular depth 3 19.5 

Distal articular length 4 22.7 

Distal articular angle 5 19° 

' The angle between the antero-medial to postero-lateral plane of the proximal tibial 

facet and the diaphyseal axis. 

1 The maximum antero-posterior diameter of the epiphysis, measured parallel to the 

talar surface. 

' The maximum antero-posterior diameter of the talar articular surface. 

4 The maximum proximo-distal diameter of the talar articular facet, measured 
parallel to the long axis of the facet. 

5 The angle in coronal plane of the talocrural articulation between the chord for the 
articular height and the diaphyseal axis. 

neck proportions. A maximum to minimum diameter index of 124.8 
for Gough's Cave 1 is relatively low (Mesolithic: 141.9 ± 13.2, N = 
4). However, the size of its neck circumference vis-a-vis midshaft 
circumference (81.3) is close to the values for other Mesolithic 
fibulae (79.8 ± 2.2, N = 4). 

Proximal Epiphysis 

The right fibula preserves a large rounded head with a subcircular flat 
facet for the tibia. It is notable primarily for its development of a very 
large ossification of the proximal tibio-fibular ligament (Fig. 11). 
The crest is rounded on its medial margin, 22.8mm long (proximo- 
distally), 13.1mm thick, and projects ca.9.0mm from the adjacent 
head. Interestingly, there is no counterpart on the proximal tibia, only 
the tapering off of the modest interosseus line previously noted. 

Distal Epiphysis 

The right distal epiphysis generally smooth in its external surfaces. 
The digital fossa is modest in size, and the malleolar surface is gently 
convex in a proximo-distal direction. The angle between the proximo- 
distal chord of the articular surface relative to the diaphyseal axis 
( 19°) fallsclose to means of variable Mesolithic (20.1° ±6.5°, N = 7) 
and Mesolithic male (21.0° ± 6.4°, N = 5) samples. 



TALUS 



Inventory 

Right (No. 1.1/29) 

A complete bone, which has had holes drilled in the medial calcaneal 

surface and the sulcus tali for analytical samples. 

Morphology 

The right talus of Gough's Cave 1 (Table 17; Fig. 15) is a modest 
bone with generally smooth surfaces. In overall length relative to 
femoral length, it is very close to other Mesolithic specimens [12.6 
versus 12.6 ± 0.4 for Mesolithic (N = 1 1 ) and male Mesolithic (N = 
6) samples]. Similarly, its relative trochlear length (versus length) 



GOUGH'S CAVE 1: STUDY OF PELVIS AND LOWER LIMBS 



17 




Fig. 14 Views of the Gough's Cave 1 right tibial epiphyses. Left: posterior proximal epiphysis and diaphysis. Above right: proximal medial 
epiphysis. Below right: anterior distal epiphysis; x 0.8 (enlargement of parts of Figs 11, 12). 



index of 65.1 is just below the means of variable Mesolithic (65.4 ± 
4.1, N = 10) and male Mesolithic (65.3 ± 4.2, N = 6) samples. 
However, its trochlea is slightly narrower than those of most 
Mesolithic tali, since a trochlear breadth/length index provides a 
value of 79.1 for Gough's Cave 1, but means of 84.5 (±7.5, N = 26) 
and 82.8 (± 6.3, N = 18) for pooled-sex and male only Mesolithic 
samples. Finally, its neck shaft angle of 23° falls in the middles of the 
ranges of Mesolithic (24. 1 ° ±4.4°, N = 7) and male Mesolithic (22.9° 
±3.5°, N = 6) samples. 

On its talo-crural articulations, it presents several variants of the 
anterior trochlear margin (Table 18). There is a full anterior exten- 
sion of the medial malleolar surface with an associated medial 
extension of the trochlear margin. The lateral trochlear margin 
likewise extends anteriorly, and abuts against a lateral squatting 
facet, the latter matching the one on the distal tibia. 

Plantarly, the medial and anterior calcaneal facets are partially 
fused. They exhibit a non-articular wedge extending 7.8mm in from 
the sulcus tali, leading up to a clear fusion line 8.4mm long. In 
addition, the posterior 8.4mm of the margin between the anterior 
calcaneal facet and the talar head along the postero-medial portion of 
their border is open. 

The bone lacks a clear sulcus tali facet on the anterior margin of 



the posterior calcaneal surface, but it presents rounding of that 
surface along its more anterior sulcus tali margin. In addition, there 
is a swelling of non-articular bone 1 4.7mm long in the antero-medial 
portion of the postero-medial end of the sulcus tali. It is up to 4.4mm 
wide, and it abuts against the posterior margin of the medial calca- 
neal facet, remaining separate from the posterior facet. 

Finally, the posterior calcaneal facet has a persistent small sulcus 
3.0mm long extending in near the posterior end of the postero-lateral 
margin of the facet. It is the result of incomplete fusion of the 
ossification center for the lateral posterior tubercle to the talus. 



CUBOID 

Inventory 

Right (No. 1.1/30). 
Complete undamaged bone. 

Morphology 

The right cuboid bone of Gough's Cave 1 (Table 19; Fig. 15) is a 



18 




Fig. 15 Dorsal (above) and plantar (below) views of the Gough's Cave 1 
right talus and cuboid bone; x 0.9. 

Table 17 Osteometries of the Gough's Cave 1 right talus. 

60.5 
55.0 
26.0 

46.7 
35.8 
28.3 
11.4 
26.5 
28.0 

9.0 
29.5 
20.0 
37.2 
21.6 
35.0 
23.5 

5° 
23° 
34° 
41° 

59° 



Maximum (lateral) length 1 
Length (M-l) 
Articular height (M-3b) 
Articular breadth (M-2b) 
Trochlear length (M-4) 
Trochlear breadth (M-5) 
Trochlear height (M-6) 
Lateral malleolar height- 
Lateral malleolar oblique height (M-7a) 
Lateral malleolar breadth (M-7) 
Lateral malleolar length 3 
Head and neck length (M-8) 
Head length (M-9) 
Head breadth (M-10) 
Posterior calcaneal length (M-12) 
Posterior calcaneal breadth (M-13) 
Trochlear angle 4 
Neck angle (M-16) 
Torsion angle (M-l 7) 
Posterior calcaneal angle (M-l 5) 
Subtalar angle 5 



1 Distance from the distal head to the proximal lateral tubercle parallel to the sagittal 
plane of the trochlea. 

2 Distance from plantar-lateral tip of the lateral malleolar surface to the highest point 
on the lateral malleolar arc, measured in the coronal and sagittal planes determined 
by the horizontal plane of the mid-trochlea. 

'The antero-posterior maximum distance on the articular surface for the lateral 
malleolus (Day & Wood, 1968). 

4 Angle between the medial and lateral margins of the middle of the trochlea 
(Trinkaus, 1975b). 

5 Angle between the long axis of the subtalar joint (midline across the medial and post- 
erior calcaneal surfaces) and the median sagittal plane of the trochlea (Trinkaus, 1975b). 





E. TRINKAUS 


Table 18 Discrete trait features of the Gough 


s Cave 1 right talus. 


Calcaneal surface fusion 


partial anterior & medial 1 


Anterior extension of the medial malleolar 


present 


surface 2 




Medial extension of the trochlea 


present 


Lateral extension of the trochlea 


present 


Medial squatting facet 


absent 


Lateral squatting facet 


present 


Sulcus tali facet 


absent 


Sulcus tali margin rounding 


present 



1 Partial fusion of the anterior and medial surfaces with a notch present along the 

sulcus tali margin (see Trinkaus. 1975a). 

2 For definitions of variations, see Bamett ( 1954) and Trinkaus (1975a). 



Table 19 Osteometries of the Gough's Cave 1 right cuboid. 

Maximum length (M-l) 
Medial length 1 
Lateral length (M-2) 
Height 2 

Calcaneal height 1 
Calcaneal breadth 4 
Navicular height 
Navicular breadth 
Lateral cuneiform height 
Lateral cuneiform breadth 
Metatarsal 4/5 height 
Metatarsal 4/5 breadth 
Metatarsal 4 height 
Metatarsal 4 breadth 
Metatarsal 5 height 
Metatarsal 5 breadth 



38.4 
29.7 
13.5 
26.6 
23.5 
30.0 
11.2 
5.8 
17.3 
13.2 
19.3 
28.5 
19.3 
12.7 
14.5 
15.8 



1 Minimum distance on the medial side between the calcaneal and metatarsal 4 
facets. 

2 Maximum dorso-plantar height of the bone. 

3 All articular facet heights are the maximum dorso-plantar dimension of the articular 
facet in question. 

4 Articular facet breadths are the maximum medio-lateral dimensions for the 
calcaneal and metatarsal facets and the maximum proximo-distal dimensions for the 
navicular and lateral cuneiform facets. 



relatively long bone that is strongly narrowed laterally. An index 
comparing its maximum length to that of the talus provides a value 
of 69.8, which is exceeded only by that of Le Peyrat 5 (70.9) in a 
small 80% male sample of Mesolithic cuboid bones (62.8 ± 5.0, N - 
5). At the same time, its index of the lateral length to maximum 
(medial) length (35.2) is the lowest of the available Mesolithic 
indices (49.7 ± 8.3, N = 5), again approached only by the one from Le 
Peyrat 5 (35.8). 

The non-articular surfaces of the bone are quite porous, and the 
articular surfaces themselves have generally distinct but rounded 
margins. There is a large facet for the navicular bone, which is 
separated from the calcaneal facet by 2.0mm of non-articular bone 
and blends into the lateral cuneiform facet with only a modest angle 
in the subchondral bone surface. The metatarsal 4 and 5 facets are 
partially separated by a vertical ridge, with the metatarsal 5 facet 
being distinctly wider but shorter. 

The peroneal sulcus exhibits a lateral projection for the tendon of 
M. pewneus longus, but its surface shows no evidence of an articu- 
lation with a sesamoid bone. 



GOUGH'S CAVE 1 : STUDY OF PELVIS AND LOWER LIMBS 



19 



METATARSALS 



Inventory 

Metatarsal 1 Right (No. 1 . 1/33). Complete bone. 
Metatarsal 3 Right (No. 1.1/32). Complete bone. 
Metatarsal 4 Right (No. 1.1/31). Complete bone. 

Morphology 

The three preserved metatarsal bones of the Gough's Cave 1 right 
foot present an unexceptional morphology, with relatively smooth 



surfaces and distinct articular facets. The metatarsal 1 presents a 
strongly twisted medial cuneiform facet and a large but smooth M. 
peroneus longus tubercle. Distally, it has a relatively flaring surface 
for the lateral sesamoid bone, and a clear lateral deviation of the head 
indicating hallux valgus. 

Robusticity indices for the Gough's Cave 1 metatarsals (geomet- 
ric mean of the midshaft diameters versus articular length) are 
similar to Mesolithic means for the first and fourth rays (metatarsal 
1 : 22.9 versus 23.0 ±1. 6, N = 5; metatarsal 4: 12.9 versus 13.1 ±0.7, 
N = 5). The third metatarsal, however, has a robusticity index (12.2) 
which is two standard deviations below the mean of a Mesolithic 
comparative sample (13.4 ± 0.6, N = 5). 




Fig. 16 Plantar (above), medial (below left) and lateral (below right) views of the Gough's Cave 1 right metatarsals; x 



20 



E. TRINKAUS 



61.7 


72.4 


71.4 


59.6 


70.4 


69.4 


14.4 


9.7 


11.3 


12.9 


7.6 


7.1 


145.9 


57.9 


63.0 


75.6 


50.7 


50.6 


70.3 


7.2 


12.4 


438.1 


331.0 


472.1 


174.9 


207.2 


193.7 


613.0 


538.2 


665.8 



Table 20 Length and midshaft diaphyseal measurements of the Gough's 
Cave 1 right metatarsals; in mm, unless otherwise noted. 



Maximum length 

Articular length (M-l, M-2) 

Midshaft height (M-4) 

Midshaft breadth (M-3) 

Total area (mm 2 ) 

Cortical area (mm 2 ) 

Medullary area (mm 2 ) 

Dorso-plantar 2nd moment of area (I x ) (mm 4 ) 

Medio-lateral 2nd moment of area (I v ) (mm 4 ) 

Polar moment of area (mm 4 ) 



Table 21 Osteometries of the Gough's Cave 1 right metatarsal epiphyses. 

1 3 4 



Proximal maximum height (M-7) 
Proximal maximum breadth (M-6) 
Proximal articular height 
Proximal articular breadth 
Lateral cuneiform breadth 1 
Dorsal metatarsal 2 height 2 
Dorsal metatarsal 2 breadth 
Plantar metatarsal 2 height 
Plantar metatarsal 2 breadth 
Metatarsal 3 height 
Metatarsal 3 breadth 
Metatarsal 4 height 
Metatarsal 4 breadth 
Metatarsal 5 height 
Metatarsal 5 breadth 
Distal height (M-9) 
Distal maximum breadth (M-8) 
Distal articular breadth 
Distal medial height 3 
Distal lateral height 
Torsion angle (M-l 1) 
Horizontal angle 4 
Vertical angle 5 
Horizontal head angle 6 



SUMMARY 



30.1 


21.0 


19.0 


21.6 


14.3 


13.1 


30.1 


20.7 


18.6 


16.8 


13.6 

8.7 
9.2 
8.4 
4.8 

11.3 
11.9 


11.9 

2.3 

10.6 
11.0 

10.3 
11.3 


22.8 


15.5 


14.4 


21.9 


10.5 


11.1 


20.2 


9.8 


9.4 


19.6 






20.9 






3° 


14° 


25° 




16° 


21° 




4° 


15° 



7° 



'Breadths of the secondary proximal metatarsal facets are all proximo-distal. 

2 Heights of the secondary proximal metatarsal facets are all dorso-plantar. 

'Distal medial and lateral heights are from each hallucal sesamoid sulci to the dorsal 

margin of the metatarsal head. 

4 Angle between the coronal plane of the main metatarsal facet and the diaphyseal 

axis in the horizontal plane of the bone. A positive angle indicates a medial deviation 

of the facet. 

5 Angle between the coronal plane of the main metatarsal facet and the diaphyseal 

axis in the sagittal plane of the bone. A positive angle indicates a plantar deviation of 

the facet. 

"Angle in the horizontal plane between the intersesamoid crest and the diaphyseal 

axis. 



The relative lengths of the Gough's Cave 1 metatarsals can be 
assessed by comparing their articular lengths to talar length and 
femoral bicondylar length. In the first comparison, the first and third 
rays produce indices of 108.4 and 128.0, which are slightly shorter 
and longer respectively than the means of a Mesolithic sample 
(metatarsal 1: 113.6 ± 7.3, N = 5; metatarsal 3: 122.9 ± 1 1.8, N = 5). 
Comparing the same lengths to femoral length produces indices of 
13.7 and 16.1, values which are similar to and slightly above the 
means of a Mesolithic sample ( 13.8 ± 0.8, N = 5 and 15.6 ± 1.1, N = 
5 respectively). 



The lower limb remains of Gough's Cave 1 are therefore those of a 
largely average young adult male, compared to other European 
Mesolithic specimens. Overall diaphyseal robusticity is generally 
similar to that of other Mesolithic specimens, even though the fibula 
and third metatarsal appear relatively gracile. In general, however, 
musculo-ligamentous attachment areas are weakly marked, in terms 
of the prominence, size and rugosity of the various crests and 
tuberosities. The exceptions to this are the marked pilasters of the 
femora (but weak lineae asperae) and the large proximal tibio-fibular 
ligament crest on the right fibula. 

The proximal femora and the femoral diaphyses exhibit a clear 
asymmetry, especially in their neck-shaft angles and diaphyseal 
dimensions. This asymmetry is accompanied, in the pelvis, by a 
greater degree of lateral flare of the left ilium. It is not possible to 
determine, given primarily preservation of only the right side below 
the knee, whether this asymmetry continued distally. 

These aspects are associated with a pelvis that combines several 
distinctly male characteristics with an overall pelvic aperture shape 
which is female. 



Acknowledgements. I would like to thank Chris Stringer for inviting 
me to participate in the Cheddar Man project. I am very grateful to Steve 
Churchill for taking over the description of the upper limb and axial remains, 
thereby relieving me of the need to sequence the ribs, and to Trent Holliday 
and Steve Churchill for dealing with issues of body size and proportions. 
Steve Churchill also collected lower limb osteometries and generated the raw 
data for the Mesolithic comparative lower limb cross sections, and Erik 
Ozolins digitized all of them. The femoral and tibial cross-sectional geometry 
samples were greatly expanded through the work of Brigitte Holt. My 
participation in this project has been supported by the Interdisciplinary 
Research Fund of the Natural History Museum (London) and National 
Science Foundation grant SBR-93 18702. To all of these individuals and 
institutions 1 am grateful. This paper was submitted and accepted for publica- 
tion in October 1997. 



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Bull. nat. Hist. Mm. Land. (Geol.) 58(supp): 23-35 



Issued 26 June 2003 



Human Dental Remains from Gough's Cave 
(Somerset, England) 



DIANE E. HAWKEY 

Department of Anthropology, Arizona State University, Tempe, AZ 85287-2402, USA 



Synopsis. The dental remains of nine individuals from Gough's Cave (Cheddar, Somerset) date from Late Pleistocene to the 
Holocene. Descriptions are provided for all individuals for crown and root morphology, odontometric data, dental pathology 
(caries, abscess, periodontal disease, enamel hypoplasia), calculus deposition, enamel pressure chipping, occlusal attrition, and 
evidence of intentional/occupational modification. The analytical focus is on seven individuals who date from the Late Upper 
Paleolithic/Mesolithic (Creswellian) culture periods. Comparative data from nine world populations suggest five trends: 1) 
Gough's Cave individuals have a morphologically simplified dental pattern similar to other Late Pleistocene/Early Holocene 
populations of North Europe, South/Southwest Asia and North Africa. 2) Within Europe, Gough's Cave is consistent in post- 
Pleistocene trend towards reduction in tooth size. 3) There is a temporal trend in the British Isles towards lateral incisor reduction, 
while maintaining stable molar tooth size. 4) Pathology, wear, and enamel pressure chipping are consistent with a hunter/gatherer 
lifeway. with one individual who may have occupationally related microtrauma. 5) No evidence occurs of any cleaning striations 
('toothpick groves') as has been suggested for Neanderthals. 



INTRODUCTION 



Little is currently known about the dentition of Late Pleistocene/ 
Early Holocene inhabitants of the British Isles. Excavations at 
Gough's Cave (Cheddar Gorge, Somerset) have recovered the dental 
remains for a minimum number of seven individuals dating to this 
time range. The remains have been radiocarbon dated to between 
1 2,380 and 9,080 BP (Hedges et al 1 99 1 ) and include 'Cheddar Man' 
(Gough's Cave 1 ), the most complete early human skeleton from 
Britain. Individuals from this time span date to the Upper Late 
Paleolithic/Mesolithic (Creswellian) culture periods. Dentition from 
two additional specimens (Gough's Cave 4 and 5) are more recent, 
dating to the Late Holocene. Gough's Cave 1 , although dating to the 
Mesolithic time period, was included in analysis of the Upper Late 
Paleolithic group in order to maximize sample. While the sample is 
small the assumption is that the available data characterize individu- 
als from early Gough's Cave. 

The first part of this study describes all dental remains from 
Gough's Cave. Included are crown and root morphology, odonto- 
metric data, pathology (caries, abscess, periodontal disease, enamel 
hypoplasia), calculus deposition, enamel pressure chipping, occlu- 
sal attrition (wear), cultural treatment and intentional/occupational 
modification. The latter half of the study focuses specifically on 



the dentition from the Late Pleistocene/Early Holocene and prov- 
ides a comparative analysis with other early and recent world 
populations. 

General descriptions of the Gough's Cave skeletal and dental 
remains have been published elsewhere (Oakley etal\91\\ Tratman 
1975; Stringer 1985, 1990). This research is part of a larger series of 
forthcoming articles published in the Bulletin of the Natural History 
Museum that will present a detailed analysis of the material. 



METHODS AND MATERIALS 

Gough's Cave remains used in this study are currently housed at the 
Natural History Museum in London and were excavated in 1903, 
1927-29, and 1986-87 (Davies 1904; Seligman & Parsons 1914; 
Keith & Cooper 1929; Cooper 1931; Currant et al 1989). Although 
Humphrey and Stringer (n.d.) argue for a numerically conservative 
approach, and suggest a minimum number of five individuals for the 
Late Pleistocene/Early Holocene group, the lack of any clear asso- 
ciation between the dental elements (especially occlusion and enamel 
pressure chipping patterns) argues for a minimum number of seven 
individuals as presented in the current study (Table 1 ). The two more 
recent specimens (Gough's Cave 4 and 5) date to the Late Holocene. 



Table 1 Gough's Cave specimen numbers, time period, age, sex, number of teeth with morphology data (includes root data and unerupted teeth), and 
number of teeth with odontometric data. 



Specimen number 



Time period 



Age 



Sex 



;y (n = teeth) 


Metrics (n = 


teeth) 


27 


18 




1 


1 




13 


12 




10 


5 




1 


1 




16 


1 




27 


20 




6 


1 




4 








87-25/87/49 


Late Pleistocene 


Adolescent 


Unknown 


87-103a 




Late Pleistocene 


Adult, mid-old 


Unknown 


87-139 




Late Pleistocene 


Adult, young-mid 


Unknown 


87-253 




Late Pleistocene 


Adult, young-mid 


Male 


89-001 




Late Pleistocene 


Adult, young 


Unknown 


Gough's 


Cave 6 


Late Pleistocene 


Adult, mid-old 


Male 


Gough's 


Cave 1 


Early Holocene 


Adult, young-mid 


Male 


Gough's 


Cave 4 


Late Holocene 


Adolescent 


Unknown 


Gough's 


Cave 5 


Late Holocene 


Adult, mid-old 


Unknown 



Total 



105 



59 



© The Natural History Museum, 2003 



24 



D.E. HAWKEY 



The dental remains described here include three males (all Late 
Pleistocene/Early Holocene), with the remaining six of indetermi- 
nate sex. The Late Pleistocene/Early Holocene adults range in dental 
age from adolescent (n = 1 ), young adult (n = 1 ), young-middle adult 
(n = 3), middle-older adult (n = 2). The two Late Holocene individu- 
als are an adolescent (Gough's Cave 4) and a middle-older age adult 
(Gough's Cave 5). Age determination for adolescents is based on 
eruption, degree of root formation, and occlusal wear. Adult age is 
based on degree of dental attrition, using the adolescent sample as a 
baseline. 

Morphology: Crown and root morphology data for 105 teeth 
were collected using the Arizona State University Dental Anthropol- 
ogy System (Turner et al 1991 ). The Dental Anthropology System 
(DAS) consists of a series of rank-scaled reference plaques to score 
trait presence and degree of expression. When congenital absence of 
a tooth was suspected, the score was confirmed through use of 
radiographs. Data for two additional crown traits (to be added to 
DAS in the future) were collected: 1 ) maxillary premolar accessory 
ridge (Burnett et al 1996) and 2) upper premolar buccal style 
(Hawkey n.d.a). 

For the purpose of analysis, the individual count was used (Turner 
& Scott 1977), a method that assumes the highest grade of expres- 
sion for a given antimere best characterizes an individual's genotype 
for that trait. Thus, the score used for an individual is the highest 
grade observed between the two sides. In order to maximize sample 
size when only one side is present, the score for that side is used, and 
symmetry is assumed. Comparative key trait data for nine geo- 
graphic populations were obtained from the literature and adjusted 
to reflect the DAS breakpoints for presence/absence following meth- 
odology used by Turner (1987). The key traits for a given tooth/ 
feature are considered to be the most reliable population discrimina- 
tors, and are scored for the teeth considered to be the least influenced 
by environmental factors according to the Field concept (Dahlberg 
1945). 

Metrics: Odontometric measurements for 59 teeth were taken 
using Helios needle-point calipers, calibrated to 0.05 mm. Each 
measurement was taken on three separate occasions and all were 
found to be within 0.05 mm difference. When discrepancies occurred, 
the results of the three measurements were averaged. The mesiodis- 
tal (MD) and buccolingual (BL) diameters of the maximum crown 
length and breadth were obtained, following the methods of Moorrees 
(1957). Teeth with observable interproximal wear were not meas- 
ured for MD diameter. In addition, data for crown height of unworn 
teeth and complete root length were collected. 

Asymmetry between right and left antimeres in both MD and 
BL diameters were assessed by paired samples t-test. A metric 
description of crown size and shape dimension was calculated by 
use of Crown Index, Crown Area, and Crown Module for all 
premolars and molars. Crown Index ([BL/MD] x 100) provides a 
measurement of relative crown breadth, with a score of 100 indi- 
cating that the BL and MD measurements are equal; a score greater 
than 100 denotes that the BL diameter is larger than the MD 
diameter. Crown Area (MDxBL) provides occlusal surface area, 
although it is assumed that the surface is rectangular. Crown Mod- 
ule ([MD+BL]/2) is calculated to indicate the average diameter of 
the tooth. While Crown Index provides some idea of occlusal 
shape, the Crown Module and Area describe the size of the crown. 
Incisor Breath (MD diameter IVMD diameter of I 1 ) was also deter- 
mined because the MD ratio of the upper incisors has been 
proposed as useful in population affinity assessment (Lukacs 1985; 
Potter et al 1981). 

When both sides were present, left side data were utilized for 
odontometric analysis. Due to the limited number of teeth available 



for analysis, right side measurements were used when the left side 
was absent (Goose 1 963 ). Data from both sexes were pooled, because 
only three males could be identified reliably in the sample. 

In order to characterize the population in a single figure, the Total 
Crown Area (sum of the mean crown area for all maxillary and 
mandibular teeth on one side) was calculated and presented as 
millimetres squared ( mm 2 ). The Molar Crown Area for M 1 -M2 teeth 
( M 1 M2C A ) was also calculated ( sum of the Crown Area of maxil- 
lary and mandibular first and second molars on one side) in order to 
assess posterior tooth size. Ordinarily, the M3 is included in the 
calculation, but a lack of M3 data in the comparative samples 
necessitated use of only M 1 and M2. The Penrose shape/size statistic 
(Penrose 1954) was used to assess dental metric population similari- 
ties based on both size and shape components; Corruccini ( 1 973 ) has 
found the shape component to be particularly useful for population 
comparisons. 

Pathology/occlusal attrition/crown chipping: Data for three 
forms of dental pathology (caries, abscessing, periodontal disease), 
calculus deposition and enamel hypoplasia were collected. Caries 
were scored for presence/location, following the definitions and 
procedures established by Koritzer (1977). An abscess was defined 
as a perforation of the alveolar bone connected to the root socket, 
while periodontal disease was noted in terms of degree of root 
exposure with antemortem erosion of the alveolar border (Turner et 
al, 1991). Calculus deposition was scored following Brothwell's 
(1981) definition of slight, medium, and heavy. Presence of enamel 
hypoplasia was scored as either chronic or acute episodes, in linear 
or pitting forms, with the dental age development estimate obtained 
from Schour and Massler's (1940) crown formation chart for U.S. 
Whites. Degree of occlusal attrition (wear) was noted for each tooth, 
following procedures established in DAS, with a score of '0' indicat- 
ing no wear, ' .5 ' as trace wear facets seen with 1 Ox magnification, ' 1 ' 
has dentine exposed, '2' indicates cusps are worn away, '3' is 
exposed pulp, and '4' as functional root stump, with all or most of the 
enamel missing. Antemortem crown chipping (microtrauma) was 
determined through examination by a lOx hand lens to differentiate 
from post-mortem damage. Any presence of chipping was noted as 
to tooth and location on the tooth. 

Other features: Any evidence of intentional dental modification 
(ablation, filing, inlay, staining), cleaning striations (brushing, inter- 
proximal 'toothpick' grooves), or occupational use of teeth were 
described. 



SPECIMEN DESCRIPTIONS: LATE 
PLEISTOCENE/EARLY HOLOCENE 



Specimen. Gough's Cave 87-25/87/49 (including Specimens 
009,120a, 120b, 165, 264b) 

Time period. Late Pleistocene (Late Upper Paleolithic) 

Description. Individual is an older adolescent (approximately 
15-18 years), based on eruption pattern and degree of occlusal wear. 
The specimen includes the left maxilla (87/25), right maxilla (87/ 
87), and mandible (87/49), including the RI, (87/264b), LI, (87/ 
102a), RP 3 (87/120b), LP 3 (87/165), and LP 4 (89/009). Maxillary 
dentition: Present are the LRI 1 2 , LR C , LP 3 , retained deciduous lm 2 , 
LM' 2 and an unerupted LM 3 . The RP 3 , RP 4 , RM 1 2 3 are missing post- 
mortem and there is postmortem damage (chipping) on the occlusal 
edges of the RI 1 2 and the paracone of LM 2 . Mandibular dentition: 
Teeth present are the RI, LI„ LRP r LP 4 , LRM, ,, and unerupted 



HUMAN DENTAL REMAINS FROM GOUGH'S CAVE 



25 



LRM,. There is postmortem loss of the LI,, RI,, LR C , and RP 4 . There 
is no evidence of cultural treatment or modification of the teeth. 

CROWN WEAR. All maxillary and mandibular anterior teeth present 
and all first molars exhibit slight to moderate (grade = 1 ) wear. All 
second molars have only wear facets present (grade = 0.5). All third 
molars are unerupted and lack wear. 



Enamel pressure chipping. 
mandibular teeth. 



None occurs on either maxillary or 



Pathology. No evidence occurs of caries, abscess, calculus, perio- 
dontal disease, or enamel hypoplasia on maxillary or mandibular 
teeth. 

Crown morphology. Maxillary dentition: LRI 1 : labial curvature 
= 1, there is no winging. LRI 1 2 : Absence of shovel, double shovel, 
interruption groove, tuberculum dentale. LRI 2 are not peg-shaped, or 
reduced. LR C : lack shovel, double shovel, tuberculum dentale, me- 
sial or distal accessory ridges. LP 3 : lacks double shovel, accessory 
cusps, disto-sagittal ridge, enamel extension, odontome, MxPAR, or 
buccal style. LM 1 : metacone = 4, hypocone = 5, lack of cusp 5. LM 2 : 
metacone = 3.5, hypocone = 3, cusp 5 = 4 with the presence of cusp 
6, (grade 1 on the UM5 plaque, with cusp 5 very much larger than 
cusp 6). Both molars lack Carabelli's trait, parastyle, and enamel 
extension. Mandibular dentition: RI, and LI,: absence of shovel. 
LRP 3 : lack enamel extension and odontome. RP,: grade = 1 lingual 
cusp. LP 4 : single lingual cusp (grade = 0). LRM,: Y-5 pattern (cusp 
5 = grade 5), protostylid = 1, enamel extension = 2, with absence of 
anterior fovea, cusp 7. LRM,: + 5 pattern (cusp 5 = grade 3), with 
absence of deflecting wrinkle, distal and mid-trigonid crest, 
protostylid, cusp 7. 

ROOT MORPHOLOGY. Maxillary dentition: LI 1 : single root, with 
one radical. LI 2 : single root with two radicals. L c : single root with 
three radicals. LP 3 : single root with two radicals. Mandibular denti- 
tion: All incisors, canines and premolars are single root teeth. RI,: 
four radicals. LI,: two radicals. RP,: presence of Tomes' root (grade 
= 3), six radicals. LP,: presence of Tomes' root, four radicals. 

Odontometric data 





crown dimensions 


crown 


root 


crown 


crown 


crown 




MD 


BL 


height 


length 


index 


area 


module 


LI 1 


9.28 


7.85 


_ 


12.20 


_ 


_ 


_ 


RI 1 


9.45 


7.73 


- 


- 


- 


- 


- 


LP 


7.38 


7.23 


- 


12.95 


- 


- 


- 


RI 2 


7.48 


7.55 


- 


- 


- 


- 


- 


L c 


7.50 


9.50 


- 


14.80 


- 


- 


- 


R c 


7.40 


9.58 


- 


- 


- 


- 


- 


LP 3 


6.43 


9.28 


- 


10.00 


144.32 


59.67 


7.86 


LM 1 


10.83 


12.83 


- 


- 


118.47 


138.95 


11.83 


LM 2 


10.23 


13.23 


- 


- 


129.33 


135.34 


11.73 


dim 2 


9.48 


11.50 


- 


- 




- 


- 


RI, 


5.70 


6.38 


- 


11.53 






_ 


LI, 


6.23 


7.03 


- 


13.48 






- 


LP, 


6.80 


8.63 


- 


11.20 


126.91 


58.68 


7.72 


RP 3 


6.78 


8.90 


- 


11.18 


131.27 


60.34 


7.84 


LP 4 


7.15 


8.50 


- 


- 


118.88 


60.78 


7.83 


LM, 


11.58 


11.15 


- 


- 


96.29 


129.12 


11.37 


RM, 


11.78 


11.10 


- 


- 


94.23 


130.76 


11.44 


LM 2 


11.90 


10.43 


- 


- 


87.65 


124.12 


11.17 


RM", 


11.20 


10.18 


- 


- 


90.89 


1 14.02 


10.69 



Specimen. Gough'sCave 87-103a 

Time period. Late Pleistocene (Late Upper Paleolithic) 

Description. An isolated LI 1 does not match any of the maxillary 



alveolar sockets present. There is perimortem damage to the distal 
third of the labial enamel surface. Occlusal wear and degree of root 
formation suggests the individual was middle-old age adult. 

Crown wear. Heavy wear occurs on the occlusal surface up to, 
but not exposing, the pulp chamber (grade = 3). An estimated loss of 
one-half of the total crown has occurred, with only the cervical one- 
half of the crown remaining. 

Enamel pressure chipping. No evidence noted of antemortem 
chipping. Postmortem chipping occurs on the distal one-third of the 
labial crown surface. 

Pathology. No carious activity is present in root or crown. No 
evidence occurs of enamel hypoplasia or calculus on the remaining 
cervical half of the crown. 

Crown morphology. Tooth is too worn to score for traits (labial 
curve, winging, shovel, double shovel, interruption groove, tubercu- 
lum dentale). 

ROOT MORPHOLOGY. LI 1 : single root, no radicals. 

Odontometric data 



crown dimensions 
MD BL 



crown 
height 



root 
length 



crown 
index 



crown 
area 



crown 
module 



LI 1 



6.98 



6.30 



Specimen. Gough's Cave 87-139 [460-B-ALT; 301.0] 

Time period. Late Pleistocene (Late Upper Paleolithic) 

Description. Specimen is of an adult maxilla, with complete 
alveolus and palate. Based on eruption and degree of occlusal 
wear, this individual is an adult, possibly young-middle age. Max- 
illary teeth present are the LRI 12 , LR C , LRP 3 , RP 4 , and LRM 1 2 , 
with antemortem loss of LP*. Postmortem damage occurred on the 
buccal surfaces of the LP 3 , the LM' and LRM 2 .The RM 3 is missing 
postmortem, and the LM 3 is congenitally absent. No evidence of 
cultural treatment or modification was found. Although Humphrey 
and Stringer (n.d.) suggest a possible association between this 
specimen and GC87-253, a lack of occlusion and lack of matching 
enamel pressure chipping patterns on the anterior dentition of 
GC87-253 argue against the two specimens representing a single 
individual. 

Crown wear. Slight-moderate, with heaviest wear on first molars 
(grade = 1.5), anterior teeth (grade = 1), and wear facets without 
dentine exposure (grade = 0.5) on the second molars. 

Enamel pressure chipping. There is a minor degree of chipping 
on the disto-labial surfaces of LRI 1 , the mesio-occlusal surfaces of 
LRI 2 , and the buccal surface of L c . 

Pathology. Antemortem loss of LP 4 has occurred. No caries, 
abscessing, or calculus noted. Slight enamel hypoplastic pitting 
(acute episode) occurs near the CEJ on LI 2 , LR C , and RM 1 . All 
remaining teeth do not display hypoplasia. 

CROWN MORPHOLOGY. LRI 1 : labial curvature = 1, tuberculum 
dentale = 1, absence of shovel, double shovel, interruption groove, 
and winging.; LRI 2 : both teeth lack shovel, double shovel, interrup- 
tion grooves, and are not peg/reduced shaped; RI 2 has tuberculum 
dentale = 2, (LP is missing data) ; LR C : absence of shovel, double 
shovel, tuberculum dentale, mesial accessory ridge; LRP 3 : no double 
shovel on RP 3 (missing data for LP 3 ), lack of mesial/distal accessory 
cusps, disto-sagittal ridge, enamel extension, odontome, buccal style 



26 



D.E. HAWKEY 



on both teeth; RP 4 : no accessory cusps, enamel extension, odontome, 
buccal style; RM 1 : metacone = 4, hypocone = 4, absence of cusp 5, 
Carabelli's trait, parastyle, enamel extension; LM': hypocone = 4, 
lack of Carabelli's trait, enamel extension; RM 2 : enamel extension = 
1 , absence of hypocone, cusp 5, Carabelli's trait; LM 2 : metacone = 4, 
hypocone = 1, absence of cusp 5, Carabelli's trait. 



Root morphology. 
Odontometric data 



Could not be determined. 



crown dimensions crown root crown crown crown 
MD BL height length index area module 



LI 1 


9.60 


7.45 


RI 1 


9.90 


7.33 


LI 2 


8.20 


7.90 


RI 2 


7.85 


7.28 


L c 


8.13 


9.85 


R c 


7.95 


9.98 


LP 3 


6.73 


9.03 


RP 3 


6.03 


- 


RP 4 


5.75 


8.83 


LM 1 


- 


12.20 


RM 1 


10.28 


- 


LM 2 


9.75 


- 



134.17 60.77 



Specimen. Gough's Cave 87-253 [304.0] 

Time period. Late Pleistocene (Late Upper Paleolithic) 

Description. Mandibular fragment of an adult male, consisting 
mainly of the right portion of the mandible, separated just distal to 
LI 2 , and including the lower portion of the right ramus. There is 
postmortem loss of LRI r RI,, RP, . The RM, is congenitally absent. 
Mandibular teeth present are the LI, (89/003) R c (87/263), RP 4 (89/ 
002), and RM ir There is no evidence of cultural treatment or 
modification. On the basis of occlusal wear and calculus deposition, 
this adult is estimated to be young-middle age. 

Crown wear. Slight-moderate, with heaviest wear (grade = 1 .5) 
on the first molar and anterior teeth (grade = 1 ), and second molars 
with wear faceting, but no dentine exposure (grade = 0.5). 

Enamel pressure chipping. None. 

Pathology. No caries, abscessing, or periodontal disease occur. 
There is possible pitting on the buccal surface near CEJ on the R c . A 
slight degree of calculus is present at the CEJ of all teeth. 

Crown morphology. LI 2 : absence of shovel; RP 4 :single lingual 
cusp (grade = 0), absence of odontomes, absence of buccal style; 
RM^ Y-5 pattern (cusp 5 = grade 4), enamel extension present (grade 
= 2), absence of protostylid, cusp 7; RM 2 : X-4 pattern, enamel 
extension present (grade = 3), absence of deflecting wrinkle, distal 
trigonid crest, mid-trigonid crest, protostylid, cusp 7. 

Root morphology. All incisors, canines and premolars are sin- 
gle-rooted; LI,: radicals = 2 ; R c : radicals = 2; RP 4 : radicals = 1. 

Odontometric data 



crown dimensions crown 
MD BL height 



root 
length 



crown 
index 



crown 
area 



crown 
module 



LL 


5.15 


6.50 


R 


7.68 


8.70 


RP, 


6.58 


8.33 


RM, 


10.08 


10.45 


RM, 


10.15 


10.58 



13.53 



126.60 54.81 
103.67 105.34 
104.24 107.39 



7.46 
10.33 
10.37 



Specimen. Gough's Cave 89-001 [Area I/Ml 02/70 1.0] 

TIME PERIOD. Late Pleistocene (Late Upper Paleolithic) 

Description. An isolated tooth, LP 4 , that does not belong to the 
same individual as Gough's Cave 89-002 or 89-003, on the basis of 
morphology, metrics or degree of wear. Amount of occlusal wear and 
degree of root formation suggest a young adult. 

Crown wear. Slight wear facets (grade = 0.5) are on the buccal/ 
occlusal surface, although no dentine is exposed. 

Enamel pressure chipping. None. 

Pathology. No caries or enamel hypoplasia, but a slight degree of 
calculus is present on the buccal and lingual surfaces of the cervical 
fourth of the crown. 

Crown morphology. Single lingual cusp (grade = 0), trace of 
buccal style (both mesial and distal), no odontome. 

Root morphology. Single rooted, with three radicals. 

Odontometric data 

crown dimensions crown root crown crown crown 
MD BL height length index area module 



LP, 



7.10 



8.25 



6.93 116.20 58.58 



100 BP [OxA-2236] 



Specimen. Gough's Cave #6 

TIME PERIOD. Late Pleistocene: 11,700 
(Late Upper Paleolithic ) 

Description. An almost complete mandible (missing right ra- 
mus) of an adult male. Only the RM, is present. There is postmortem 
loss of LRI, „ LR r , LRP, „, LRM, , and LM,. Both LRM, are congeni- 

1.2 C 3.4' 1 2 3 ° 

tally absent. There is no evidence of cultural treatment or modification 
on the remaining tooth. The individual appears to be middle-old age 
on the basis of occlusal wear and amount of root exposure. 



Crown wear. 
RM,. 



Moderate wear (grade = 2) of dentine exposure on 



Enamel pressure chipping. None. 

Pathology. No evidence for abscessing occurs. No caries, calcu- 
lus, enamel hypoplasia present on remaining tooth, although there is 
a slight degree (1-2 mm) of root exposure. 



Crown morphology. 

7. 



RM,: 4-cusped, lacking a protostylid, cusp 



Root morphology. All incisors, canines and premolars are sin- 
gle-rooted. Both first molars are two-rooted, although the root 
sockets of RM, indicate that the mesial root is slightly bifurcated 
(approximately a fourth of the total root length). The RM 2 is three- 
rooted and the socket for LM, has a mesial and distal root socket with 
a small auxiliary lingual root socket positioned just distal to the 
mesial alveolar socket, suggesting a 3-rooted tooth. 

Odontometric data 

crown dimensions crown root crown crown crown 
MD BL height length index area module 



RM, 



12.03 



10.90 



90.61 131.13 



11.47 



Specimen. Gough's Cave #1 ['Cheddar Man'] 

TIME PERIOD. Early Holocene: 9,080 ±1 50 BP[BM-525]; 9, 100: 
100 BP [OxA-814] (Mesolithic/Creswellian) 



HUMAN DENTAL REMAINS FROM GOUGH'S CAVE 



27 



Description. Adult male with an almost complete mandible (miss- 
ing left coronoid process and the left and right condyle) and an 
almost complete maxilla (missing the palate). Maxillary teeth: Present 
are the LRM 1 2 -' with postmortem loss of RI 1 2 , R c , RP\ LRP 4 . The 
remainder of the teeth (LI 1 - 2 , L c , LP 1 ) were damaged postmortem and 
observations were not made. Mandibular teeth: The LRI , LR , 
LP, ., and LRM, , 3 are present. Postmortem loss of RP, 4 had occurred. 
There was no evidence of any cultural treatment or modification of 
the teeth. On the basis of eruption and occlusal wear, the individual 
was of young-middle age. 

Crown wear. Moderate to slight, with heaviest wear on LP 4 , and 
RM , (grade =1.5), remaining anterior teeth and LM (grade = 1 ), and 
all remaining molars (grade = 0.5). 

Enamel pressure chipping. RM 1 : lingual portion of the hypo- 
cone. 

Pathology. No evidence occurs of caries, abscessing, periodontal 
disease, enamel hypoplasia, or calculus. 

Crown morphology . Maxillary teeth : LRM ' : metacone = grade 
4, hypocone = grade 4, absence of cusp 5, Carabelli's trait, parastyle, 
and enamel extension. RM 2 : metacone = 4, hypocone = 3, cusp 5 = 
3, with lack of Carabelli's trait, parastyle, enamel extension. LM 2 : 
metacone = 4, hypocone = 1 , with absence of cusp 5, Carabelli's trait, 
parastyle, and enamel extension. LRM 1 : metacone = 3.5, hypocone 
= 1, with absence of cusp 5, Carabelli's trait, parastyle, enamel 
extension. Neither third molar exhibited a peg or reduced form. 
Mandibular teeth: All incisors lacked shovel. L c : absence of distal 
accessory ridge. LP,: lacked a lingual cusp (grade = A), and both 
LP, did not have enamel extension, or odontomes. LRM,: have the 
Y-5 pattern (cusp 5 grade = 5), and lack protostylid, cusp 7, and 
enamel extension. LRM, and LM 3 :have an X-4 pattern, with absence 
of deflecting wrinkle, distal and mid-trigonid crests, protostylid, 
cusp 7, and enamel extension. RM,: has a Y-4 pattern and also lacks 
deflecting wrinkle, distal and mid-trigonid crest, protostylid, cusp 7. 
Because molars are slightly crowded, the torso-molar angle could 
not be assessed. 

Root morphology. RP, 4 : single-rooted. 
Odontometry data 





crown dimensions 


crown 


root crown 


crown 


crown 




MD 


BL 


height 


length index 


area 


module 


LM 1 


10.80 


11.53 


_ 


106.76 


124.52 


11.17 


RM 1 


10.95 


11.55 


- 


105.48 


126.47 


11.25 


LM 2 


10.00 


11.68 


7.18 


116.80 


116.80 


10.84 


RM 2 


10.95 


11.75 


7.58 


107.31 


128.66 


11.35 


LM 3 


8.33 


11.13 


7.50 


133.61 


92.71 


9.73 


RM 3 


9.20 


11.38 


7.88 


123.70 


104.70 


10.29 


LI , 


4.65 


5.60 


- 


- 


- 


- 


W , 


4.93 


5.55 


- 


- 


- 


- 


u, 


5.15 


6.08 


- 


- 


- 


- 


RL, 


5.25 


5.93 


- 


- 


- 


- 


L r 


6.05 


7.63 


- 


- 


- 


- 


R r 


6.48 


7.50 


- 


- 


- 


- 


LP, 


6.40 


7.33 


- 


114.53 


46.91 


6.87 


LP, 


6.83 


7.98 


- 


116.84 


54.50 


7.41 


LM, 


11.03 


10.28 


- 


93.20 


113.39 


10.66 


RM, 


11.18 


10.33 


- 


92.40 


115.49 


10.76 


LM, 


10.03 


9.80 


8.13 


97.71 


98.29 


9.92 


RM, 


10.03 


10.08 


7.20 


100.50 


101.10 


10.06 


LM, 


10.68 


10.00 


8.18 


93.63 


106.80 


10.34 


RM, 


10.83 


10.03 


- 


92.61 


108.62 


10.43 



SPECIMEN DESCRIPTIONS: LATE 
HOLOCENE 



Specimen. Gough's Cave #4 [Cooper 1929 No. 7] 

Time period. Late Holocene 

Description. Maxillary fragment separated along mid-line of a 
young-late adolescent individual. Age category was made on the 
basis of dental eruption, root formation and occlusal wear. The LM 1 
is present, and the crown of the LM 3 is complete although unerupted, 
with the roots not yet formed. Postmortem loss occurred of LI 1 2 , L c , 
LP 34 , LM 2 . Alveolar sockets indicate the roots of LP 34 are com- 
pletely formed, while the roots for LM 2 have not completely formed. 
There is no indication of cultural modification or treatment of the 
teeth. 

Crown wear. The LM' has slight wear faceting with no dentine 
exposure (grade = 0.5). 

Enamel pressure chipping. None. 

Pathology. No caries, abscessing, periodontal disease, or enamel 
hypoplasia present, although there is slight-moderate formation of 
calculus on the buccal surface near CEJ. 

Crown morphology. LM 1 : metacone = 4, hypocone = 3.5, 
Carabelli's trait = 2, absence of cusp 5, and enamel extension. There 
is a very faint parastyle expression, although it is not the buccal pit 
form (grade 1 ) and the expression of the trait is too weak to classify 
it as grade 2, because it lacks a free apex. 

Root morphology. LI 1 2 , L c , LP 3 ' 4 : single-rooted teeth. 

Odontometry data 

crown dimensions crown root crown crown crown 
MD BL height length index area module 



LM 1 



10.10 



5.85 



97.82 99.79 



9.99 



Specimen. Gough's Cave #5 [Cooper 1929] 

Time period. Late Holocene 

Description. Right mandibular fragment of an adult, with post- 
mortem loss of R c , RP, 4 , RM,. The RM, is present. Degree of 
occlusal wear suggests middle-old age. 

Crown wear. The RM, is heavily worn, with the pulp chamber 
almost exposed (grade = 2.5). 

Enamel pressure chipping. RM,:buccal and mesio-buccal sur- 
faces. 

Pathology. No caries, abscessing, periodontal disease, or enamel 
hypoplasia, although a slight-moderate degree of calculus is present. 

Crown morphology. RM,:absence of enamel extension. 

Root morphology. R c , RP, 4 : single-rooted teeth. 

Odontometry data. None available due to wear. 



28 



D.E. HAWKEY 



Table 2 Number of Gough's Cave individuals (Late Pleistocene/Early 
Holocene) with any morphological expression for a given trait. 

Data are presented only for grades that were present in the sample. Traits 
lacking any morphological expression in the sample are winging (UI1 ), labial 
curvature (UI1), shovel (UI1-2, UC, LI1-2), double-shovel (UI1-2, UC, 
UP3), interruption groove (UI1-2), canine mesial accessory ridge (UC), 
canine distal accessory ridge (UC, LC), premolar distal accessory cusps 
(UP3-4), maxillary premolar accessory ridge (UP3 only), premolar buccal 
style (UP3-4), disto-sagittal ridge (UP3), odontomes (UP3-4, LP3-4), 
Carabelli's trait (UM1-2-3), parastyle (UM1-2-3), anterior fovea (UM1), 
deflecting wrinkle (LM2-3 only), distal trigonid crest (LM2-3 only), cusp 6 
(LM1-2-3), cusp 7 (LM1-2-3), mid-trigonid crest (LM2-3 only). 

All upper and lower incisors, canines and premolars are single-rooted. The 
only data available for molars are from a single individual who has both lower 
Ml (two-rooted) and a lower M2 (three-rooted). 

Tuberculum Dentale 



grade 


Ull 


U12 UC 



1 

2 


1 
1 



1 2 



1 2 


Total 


2 


2 4 


Upper 


Molar 


Cusp 5 


grade 


UM1 


UM2 UM3 




3 
4 


3 




1 1 
1 
1 


Total 


3 


3 1 



Metacone 




grade UM1 


UM2 UM3 


3.5 
4 3 


1 1 

2 


Total 3 


3 1 



Hypocone 






grade 


UM1 


UM2 


UM3 


1 





1 


1 


3 





2 





4 


2 








5 


1 








Total 


3 


3 


1 



Enamel Extension 



grade UP3 UP4 UM1 UM2 UM3 




2 13 2 1 
10 10 




Total 2 13 3 1 




Peg, Reduced, Congenital Absence 




grade UI2 UP4 UM3 LI1 LP4 LM3 


+ 322562 
10 1 


Total 3 2 3 5 6 3 



Lower Molar Cusp Pattern 



grade LM1 


LM2 LM3 


Y 3 
X 
+ 


1 
2 

1 


Total 3 


3 1 



Lower Molar Cusp Numb er 



grade 


LM1 


LM2 


LM3 


4 
5 




3 


3 
1 


1 



Total 


3 


4 


1 



Premolar Lingual Cusp 


grade LP3 LP4 




2 
A 1 

1 1 




Total 2 2 




Lower Molar Cusp 


5 


grade LM1 LM2 LM3 


4 3 1 

5 3 10 


Total 3 4 1 



Protostylid 




grade LM1 


LM2 LM3 


2 

1 1 


4 1 



Total 3 


4 1 



Tomes 


Root 


grade 


LP3 


1 
3 1 


Total 


2 



RESULTS: LATE PLEISTOCENE/EARLY 
HOLOCENE REMAINS 

Morphology. Grades for teeth with a given trait (number or indi- 
viduals = 7, number of teeth = 97) are summarized in Table 2, and 
include root number data obtained from the alveolar socket when 
postmortem loss of a tooth occurred. In general, Gough's Cave 
individuals have a simplified, modern Homo sapiens dental pattern, 
and lack strong expression for almost all traits. 

Presence/absence breakpoints for key tooth/trait combinations 
limit the sample available for analysis due to use of the individual 
count (number of key trait observations = 65). There is a lack of 
winging (0/2), shovel I' (0/2), double shovel I 1 (0/2), interruption 
groove I 2 (0/2), upper canine mesial accessory ridge, or 'Bushman's 
canine' (0/2), upper canine distal accessory ridge (0/1 ), disto-sagittal 
ridge or 'Uto-Aztecan premolar' P 3 (0/2), Carabelli's trait M 1 (0/3), 
cusp 5 M 1 (0/3), parastyle M 3 (0/1), enamel extension M 1 (0/4), 
greater than one lingual cusp P 4 (0/2), Y-groove M (0/3), cusp 6 M [ 
(0/3), cusp 7 M (0/3), two-rooted lower canine (0/4), three-rooted 
lower molar (0/1), one-rooted M, (0/1), or upper/lower premolar 
odontomes (0/5). There are some instances of presence of tubercu- 
lum dentale I 2 (1/2), peg/reduced/congenital absence of M 3 (1/3), 
protostylid M 1 (1/3), Tomes' root P (1/2). Higher frequencies were 
noted for presence of one-rooted P 3 (2/2), presence of hypocone M 2 
(2/3) and four-cusped M, (3/4). 

Metrics: Out of 58 teeth measured in the Late Pleistocene/ 
Early Holocene sample, a total of 16 permanent maxillary teeth 
and 21 permanent mandibular teeth supplied data for the seven 
individuals, with the means summarized in Table 3. Fluctuating 
dental asymmetry in tooth size is present in the sample, although 
the results from paired t-tests between antimeres do not indicate 
significant differences statistically (p = 0.05). Only one individual 
(Gough's Cave 6) had all maxillary or mandibular molar teeth 
present. Crown Area in Gough's Cave 6 indicates an upper molar 
decrease from M 1 > M 2 > M 3 . Lower molars follow a slightly 
different pattern of M, > M, < M r 

The TCA for the Gough's Cave sample is 1,244 mm 2 (I1-M3) and 
1 ,034 mm 2 (II -M2), while the M 1 M2CA is 486 mm 2 . There is a lack 
of reduction in lateral incisor MD diameters when compared to the 
MD measurement for the central incisors, with the incisor breadth 
(I 1 : I 2 ) ratio of 0.83. 

Pathology/occlusal attrition/crown chipping: All seven indi- 
viduals could be assessed for pathology, although of these, three 
were represented by isolated teeth. There is an apparent lack of 
caries and abscessing in this series. Antemortem tooth loss (LP 4 ) 
occurred in only one individual (87/139). This same young adult 
also has the only instance of macroscopically observable enamel 
hypoplasia, an acute episode with pitting which occurs at the 
cemento-enamel junction (CEJ) of three teeth (I 1 , c , M 1 ) during 
crown formation between three to five years of age. Although no 
periodontal pockets occur, one adult male (Gough's Cave 6) showed 
evidence of 1-2 mm of root exposure of M„ the only tooth remain- 
ing, the rest having been lost postmortem. There does seem to be 
higher degree of calculus deposition (slight grades: confined to 
crown but not extending to the CEJ) occurring in two young-middle 
age adults (87-253, 89-001), including only one of the three identi- 
fiable males in the sample. 

The degree of attrition in the sample is similar to other hunter/ 
gatherer groups, in that the rate of wear is not excessive (i.e., the pulp 
chamber is not exposed before secondary dentine is formed to 
protect tooth integrity). Thus, the majority of wear seems likely to be 
age-related rather than occupational or due to highly abrasive diet. In 



HUMAN DENTAL REMAINS FROM GOUGH'S CAVE 



29 



Table 3 Summarized odontometric data for Gough's Cave sample (Late 
Pleistocene/Early Holocene), including number of individuals fn). mean 
mesio-distal diameter (MD), mean bucco-lingual diameter (BL). 
standard deviation (sd) and mean crown area. Data are for the left side, 
with right side substituted when the left is missing. When only one tooth 
is available, data are presented in parentheses. 



COMPARATIVE ANALYSIS 



Tooth 


n 


Avg. 
MD 


sd 


n 


Avg. 
BL 


sd 


Crown 
Area 


Ull 


2 


9.44 


0.23 


3 


7.43 


0.44 


70.14 


UI2 


2 


7.79 


0.58 


2 


7.57 


0.47 


58.97 


UC 


2 


7.82 


0.45 


2 


9.68 


0.25 


75.70 


UP3 


2 


6.58 


0.21 


2 


9.16 


0.17 


60.27 


UP4 


1 


(5.75) 


- 


1 


(8.83) 


- 


(50.77) 


UM1 


3 


10.64 


0.31 


3 


12.19 


0.65 


129.70 


UM2 


3 


9.99 


0.24 


2 


12.46 


1.10 


124.48 


UM3 


1 


(8.33) 


- 


1 


(11.13) 


- 


92.71 


LI1 


2 


5.18 


0.74 


2 


5.99 


0.55 


31.03 


LI2 


3 


5.51 


0.62 


3 


6.54 


0.48 


36.04 


LC 


2 


6.87 


1.15 


2 


8.10 


0.85 


55.65 


LP3 


2 


6.60 


0.28 


2 


7.98 


0.92 


52.67 


LP4 


4 


6.92 


0.26 


4 


8.27 


0.22 


57.23 


LM1 


3 


10.90 


0.76 


3 


10.64 


0.44 


115.98 


LM2 


4 


11.03 


1.09 


4 


10.50 


0.34 


115.82 


LM3 


1 


(10.68) 


- 


1 


(10.93) 


- 


(116.73) 



adults, the wear on the anterior teeth is almost always grade 1 (some 
dentine exposed). There is one instance of an isolated upper incisor 
(87- 103a) that is heavily worn (grade 3) although probably due to 
age-related wear consistent with older age. The maxillary first 
molars have a wear range of grades 1-1.5, with the lower molars 
experiencing slightly more wear (grades 1-2.5), although again, the 
type of wear suggests older adults. All erupted maxillary and man- 
dibular second and third molars in the sample were only slightly 
worn (grade = 0.5). 

The degree of wear may be correlated with instance of molar 
crown pressure chipping in at least one individual. In Gough's 
Cave 1, the RM 1 , has an attrition score of grade 1, although the 
tooth exhibits chipping along the lingual surface of the hypocone; 
its antimere, LM 1 , has the same amount of wear but lacks any 
evidence of pressure chipping. However, the mandibular counter- 
parts to these two teeth indicate much heavier use of the right side, 
with the RM! wear grade of 1.5, (no chipping) while the LM, 
mirrors the lesser wear (no chipping) of the left maxillary molar. 
Two other individuals (87-253, Gough's Cave 6), have a high 
degree of attrition on the lower molars, but wear is not correlated 
with pressure chipping. 

Specimen 87- 1 39, a young-middle age adult, displays an attrition/ 
chipping pattern that may be consistent with occupational use of the 
teeth. Crown microtrauma occurs on the maxillary incisors, between 
the right I 1 - 2 , and the left I 1 ' 2 , and on the buccal surface of the left 
upper canine. The LP 4 was lost antemortem, and the LP 3 crown was 
damaged postmortem. The left side of the anterior dentition may 
have been utilized more in this adult; the R c , RP 34 , and LRM 1 2 did 
not display evidence of chipping. Because there are no instances of 
caries in the sample, the antemortem loss of LP 4 may be related to the 
degree and amount of pressure chipping found in the dentition of this 
specimen. Unfortunately, no mandibular dental remains for this 
adult were recovered. 

Other Features: There is no evidence of cultural treatment (i.e., 
interproximal 'toothpick' grooves, enamel cleaning striations) or of 
intentional dental modification. 



Results of this comparative analysis are based on a numerically 
limited series should be treated with caution. The analysis presented 
here suggests only possible trends present in the available data, and 
with the assumption that these few individuals from Gough's Cave 
are representative of the populations of the Late Pleistocene/Early 
Holocene British Isles. 

Morphology: Table 4 presents a comparison of the Gough's 
Cave morphological data with the occurrence of 20 key traits in 
seven early populations: Upper Paleolithic-Neolithic North Europe 
(ca. 32,000-4,000 BP), Mesolithic Nubia (ca. 18,000-12,000 BP), 
Iberomaurusian North Africa (ca. 16,700-10,500 BP), Epi- 
Paleolithic Levant Natufians {ca. 12,800-10,200 BP), 
Mesolithic-Neolithic South Asia (ca. 8,000 -2,800 BP), Neolithic- 
Bronze Age Lake Baikal (ca. 7,400-3,800 BP), and Jomon Japan 
(ca. 7,000-2,300 BP). The data are also compared with two historic 
pooled populations: North Europe, and Khoisan-speakers of sub- 
Saharan Africa. Results were compared with the Early World Average 
for each trait (Hawkey 1998) . Populations were then designated as 
having percentages higher than (H), lower than (L) or within five 
percent above or below the Early World Average (A). Because the 
Gough's Cave sample is so small numerically, the frequency of each 
trait is characterized as having total absence (0), less than or equal to 
50 percent (+), or more than half the sample (++). 

Among world populations, the Gough's Cave dental morphology 
seems most like other early world groups with a simplified dental 
pattern. Gough's Cave is most similar to Late Pleistocene/Late 
Holocene (Neolithic) North Europe, sharing trait frequencies of 
83.3%. By historic times, Gough's Cave is still like Recent North 
Europe (72.2%), although there are differences in presence of tuber- 
culum dentale, hypocone, Carabelli's trait, one-root P 3 , and cusp 7. 
The simplified dental pattern seen in Gough's Cave shares similari- 
ties with two other Late Pleistocene/Early Holocene world samples: 
Epi-Paleolithic Levant Natufians (70.0%) and North African 
Iberomaurusians (70.0%), and with the Early Holocene sites of 
South Asia (83.3 %) and Jomon (80.0%). 

The sample is dentally unlike the Late Pleistocene sites of Nubia 
(42.1%), and the Late Holocene (Neolithic-Bronze Age) inhabitants 
of Lake Baikal (52.6%). If the Khoisan-speaking populations from 
modern sub-Saharan Africa are taken to represent the early sub- 
Saharan African dental pattern (Irish 1993, 1998), then Gough's 
Cave shares only 60% of key traits with this geographic group. 

Only one archaic dental trait appears in the sample. Presence of P 
Tomes' root occurred in one out of two individuals who could be 
scored for the trait. A study of more than 7,700 individuals (Turner & 
Hawkey 1991) found that the trait occurs most often in Africa, 
ranging from 20-50%, with the Late Pleistocene Nubian sample 
among the highest in the world (47%). North Europe averages 9%of 
trait presence, although the range can be quite broad (0^43%). The 
instance of Tomes' root in this sample, therefore, cannot exclude 
similarities with either European or African populations for the trait. 
However, the lack of expression in Gough's Cave of many other 
archaic traits found by Irish (1993, 1998) in the sub-Saharan African 
Dental Complex (i.e, high frequencies of upper canine mesial acces- 
sory ridge, Carabelli's trait M 1 , cusp 7 M r presence of Y-groove 
pattern M,, two-rooted P\ and lack of congenital absence M 3 ) 
suggests that Gough's Cave is dentally unlike the African samples. 

Surprisingly, Gough's Cave lacked any expression of Carabelli's 
trait (0/3), although the trait is often found to occur in high frequen- 
cies (80%) among modern British (Goose & Lee 1971). The trait is 
of little use by itself to differentiate at geographic population levels 



30 



D.E. HAWKEY 



Table 4 Results for 20 dental traits for Gough's Cave, seven Early populations, and two Recent populations (no. of individuals = 296-1494), compared 
with Early World average (based on nine geographic populations (North Asia, Nubia, Southeast Asia, Malaysia, South Asia, Early Eurasia, North Europe, 
Levant, and North American Paleo-Indian). Data for the Early World Average are taken from personal observations and literature (as cited in Hawkey 
1998). 

= absence of any expression; + = some expression (less than or equal to 50% of the sample); ++ = more than half the sample; L = less than 5% of the 
Early World Average ; A = within 5% more than or less than the Early World Average; H = more than 5% of the Early World Average. Similarities in trait 
expression percentage indicated in boldface. Gough's Cave score of "0" is similar to "0" and "L", while "+" is similar to "L" or "A". Scores of "++" would 
be equal to either "A" or "H". 











Early 










Early 




Early 


Trait 


Gough's 


Early 


Early 


South 


Early 


Early 


Recent 


Early 


North 


Recent 


World 




Cave 1 


Jomon 2 


Baikal 3 


Asia 4 


Levant 5 


Europe 6 


Europe 7 


Nubia 8 


Africa" 


Khoisan 10 


Average 


Wing 





A 


H 


L 








L 


H 


L 


A 


17.9% 


Shov 





A 


H 


H 


L 


A 


L 


H 


A 


H 


28.2% 


DbShov 





L 


H 


L 


L 


A 


L 





L 





19.5% 


IG 





H 


H 


H 


L 





— 


L 


H 


L 


27.7% 


TD 


+ 


L 


L 


L 


H 


H 


H 


L 


L 


A 


48.0% 


MAR 





L 


A 





H 


A 





H 


A 


H 


7.7% 


Hypo 


++ 


A 


H 


H 


H 


A 


L 


A 


H 


L 


88.7% 


UMC5 





A 


— 


A 


L 


L 


L 


A 


A 


A 


26.1% 


Para 





A 


H 


A 


A 





A 





A 


A 


3.4% 


Cara 





L 


A 


A 


L 


L 


H 


H 


H 


H 


28.3% 


PRCA 


+ 


A 


A 


L 


H 


— 


— 


— 


L 


H 


16.3% 


EnExt 





A 


A 











L 


H 


L 





14.4% 


1RUP1 


++ 


H 


H 


— 


L 


— 


L 


L 


L 


H 


56.3% 


PLC# 





A 


A 


L 


A 


H 


L 


H 


H 


A 


67.4% 


Y 





H 


L 


H 


H 


L 


L 


H 


H 


H 


20.9% 


6CLM1 





H 


H 


L 


L 


L 


L 


A 


L 


L 


26.0% 


4CLM2 


++ 


L 


L 


H 


L 


H 


H 





L 


L 


71.4% 


Proto 


+ 


L 


H 


L 


L 


H 


L 


H 


A 


A 


22.6% 


C7 





A 


H 


A 


A 


L 


H 


A 


A 


H 


6.8% 


Tomes 


+ 


L 


A 


— 


A 


L 


L 


H 


L 


A 


21.3% 


Totals 




16/20 


10/19 


15/18 


14/20 


15/18 


13/18 


8/19 


14/20 


12/20 








80.0% 


52.6% 


83.3% 


70.0% 


83.3% 


72.2% 


42.1% 


70.0% 


60.0% 





1 Present study 

2 Turner 1987 
'Turner 1987 

4 Hawkey 1977; Lukacs 1986 

5 Lipschultz 1996 

6 Alexandersen 1963; Beillard, cited in Brabant 1976; Brabant 1971, 1976; Brabant & Ketelbant 1975; Brabant et al. 1961;deTerra 1905; Haeussler 1996; Hellman 1928; Turner 

6 Benjamin 1989; Turner & Hawkey 1991, 1998 

7 Axelsson & Kirveskari 1977; Brabant & Ketelbant 1975; Goose & Lee 1971; Guigui 1974; Hjelmman 1928; Kaczmarek 1981; Kirveskari 1974; Lavelle et al. 1970; Morris 
1975; Morris et al. 1978; Pedersen 1949; Sauter & Moeschler 1960; Schwerz 1917; Scott 1973, 1977; Selmer-Olsen 1949; Turner & Benjamin 1989; Turner & Hawkey 1991, 
1998; Zubov & Kaldiva 1979 

8 Irish 1993 
» Irish 2000 
'"Irish 1993 



(Turner & Hawkey 1998) however, with the range in modern North 
Europeans (29-80%) quite variable in terms of trait presence. 
Although presence of Carabelli's trait averages 43% in modern 
North Europe, the same percentage is found in North American 
Indians, and is often even higher (53%) in modern sub-Saharan and 
West Africans. 

Thus, by the Late Pleistocene/Early Holocene, Gough's Cave 
shows the strongest dental similarity with Noth Europe, South/ 
Southwest Asia and North Africa. They appear to be dentally unlike 
either East Africa (Nubia), South Africa (modem Khoisan), or North 
Asia (Baikal). 

Metrics: While the amount of environmental influence on tooth 
size is debatable (Goose 1963; Hillson 1986; Kieser 1990; Lukacs 
1985; Scott & Turner 1988, 1999), crown shape appears to be 
another reliable way to assess population affinity (Corruccini 1973). 
When compared with TCA results for earlier samples from Europe 
(Table 5) Gough's Cave is approximately 13% smaller than Euro- 
pean Neanderthal dentition (based on I1-M3), but only 3% smaller 



than Late Upper Paleolithic Europeans. Gough's Cave is closest to 
other European Mesolithic populations, and have teeth 4% larger 
than Neolithic Europeans, and 10% larger than modern Europeans. 
The latter result supports the post-Pleistocene trend towards dental 
reduction, most likely due to relaxed selection for large tooth/body 
size (Brace & Mahler 1971; Wolpoff 1971). 

Table 5 Temporal comparisons of Total Crown Area (TCA) for early to 
recent Europe. Gough's Cave value is calculated with M3 data to 
compare with published information. (Source for non-Gough's Cave 
sample; Brace et al 1991). 



TCA (mm 


2 )I1-M3 


Period 


1415 




Neanderthal 


1267 




Late Upper Paleolithic Europe 


1237 




Mesolithic Europe 


1244 




Gough's Cave (Late Upper Paleolithic-Mesolithic) 


1196 




Neolithic Europe 


1127 




Modern Europe 



HUMAN DENTAL REMAINS FROM GOUGH'S CAVE 



31 



Table 6 Total Crown Area (TCA) for Gough's Cave (Late Pleistocene/Early Holocene) compared to early and modem populations. The TCA is calculated 
without M3 data. Samples that contain only males are indicated as M; pooled data for both sexes are designated as M&F. 



TCA (mm 2 )Il-M2 


Area/Site 


Sex 


Source 


1158 


Nubia (Mesolithic East Africa) 


M 


Calcagno 1986 


1103 


Mahadaha (Mesolithic India) 


M&F 


Lukacs & Hemphill 1992 


1054 


Natufian (Epi-Paleolithic Levant) 


M&F 


Dahlberg 1960 


1037 


Mehrgarh (Neolithic Pakistan) 


M&F 


Lukacs 1985 


1034 


Gough's Cave (Late Paleolithic-Mesolithic) 


M&F 


Present study 


981 


Jomon (Early Japan) 


M 


Brace & Nagai 1982 


981 


Anglo-Saxon (Early Britain) 


M 


Lavelle 1968 


966 


Britain (Recent) 


M 


Lavelle 1968 


910 


Khoisan (Recent South Africa) 


M 


Haeussler et al 1989; van Reenan 1982 



In order to compare Gough's Cave with published data for 
Holocene populations, the results are based on measurements for 1 1 - 
M2 (Table 6). According to Brace (1980), differences of more than 
100 mm 2 summed TCA are significant statistically, while differences 
of more than 50 mm 2 are probably significant. The TCA values for 
Gough's Cave are closest with the incipient agriculturalists of 
Neolithic Mehrgarh (Pakistan) with only 3 mm 2 difference, and 
Levant Natufians (20 mm 2 ). Four other populations were within 100 
mm 2 difference: the Anglo-Saxons (53 mm 2 ), Jomon (53 mm 2 ), 
Recent Britain (68 mm 2 ) and Mesolithic Mahadaha in Indo-Gangetic 
India (69 mm 2 ). Gough's Cave TCA values are unlike those of Late 
Pleistocene Nubia (124 mm 2 ) and modern sub-Saharan Khoisan- 
speakers (124 mm 2 ). 

A similar pattern appears in the Penrose size component (Table 7), 
with Levant Natufians (0.02), Mehrgarh (0.04) and Mahadaha (0.05) 
most similar to Gough's Cave. The component for Anglo-Saxon 
(0.33), Jomon (0.39), and Nubia (0.59) show less similarity. Recent 
Britain (0.97) and Khoisan (1.14) are least like Gough's Cave in 
occlusal crown size. The two groups that show closest similarity in 
Penrose shape are the Natufians (0.87) and Mehrgarh (0.97), mirror- 
ing the size component results. However, the remaining sample 
indicates moderate similarity (Jomon = 1.07,Nubia= 1.10,Mahadaha 
= 1 .24, Khoisan = 1 .34) with Anglo-Saxon (2.39) and Recent Britain 
(4.22) the most dissimilar to Gough's Cave in shape component. 

The differences between the size and shape results may be due to 
several factors. The most likely explanation is that the size compo- 
nent results reflect sexual dimorphism in tooth size. Gough's Cave, 
Natufians, Mehrgarh and Mahadaha (the most similar in size compo- 
nent results) are all from pooled samples of males and females. All 
other data are from males only. 



Table 7 Results of Penrose statistic (Shape/Size) for Gough's Cave 
sample (Late Pleistocene/Early Holocene) compared with eight other 
populations. 





Gough' 


s Cave sample 






Size 


Shape 


Combined 




component 


component 


statistic 


Natufian (M&F) 


0.02 


0.87 


0.89 


Mehrgarh (M&F) 


0.04 


0.97 


1.01 


Mahadaha (M&F) 


0.05 


1.24 


1.29 


Anglo-Saxon (M) 


0.33 


2.39 


2.72 


Jomon (M) 


0.39 


1.07 


1.46 


Nubia (M) 


0.59 


1.10 


1.69 


Recent British (M) 


0.97 


4.22 


5.19 


Khoisan (M) 


1.14 


1.34 


2.48 



Another possibility is that the TCA and Penrose size statistic, both 
of which include anterior and posterior teeth, may reflect apportion- 
ment differences within populations (Harris & Rathbun 1991). Size 
differences between incisor/canine and premolar/molar fields within 
individuals may explain why Recent Britain appears so dissimilar to 
Gough's Cave in size component, yet so similar in TCA value. 
Incisor breadth ratio (Table 8) shows that Recent Britain has the most 
reduced lateral incisors (IB = 0.72) when compared to Gough's Cave 
(IB = 0.83), yet the molar crown area for M1M2 (Table 9) indicates 
the molars are similar in size for both groups. 

When both metric and morphology differences are compared 
(Table 10) the data reveal the following consistent patterns: 

1 . Gough's Cave is most like early populations of South/Southwest 
Asia, including Pakistan (Mehrgarh) and the Levant (Natufians) 
in TCA and Penrose size/shape components. 

2. Gough's Cave is also similar to other early North Europe popul- 
ations (in TCA and DAS), although published data for North 
Europe were unavailable for Penrose size/shape analysis. 

3 . Both metric and morphology results suggest that Late Pleistocene 
East Africa (Nubia) and sub-Saharan Africans (modern Khoisan) 
are most dissimilar to Gough's Cave. The DAS morphological 
data for Late Pleistocene North Africans (Iberomaurusian) sug- 
gest a much closer dental similarity to Gough's Cave than other 
African regions. 

4. Differences within the British Isles suggest Gough's Cave is 
unlike Anglo-Saxon (53 mm 2 difference) and Recent Britain (68 
mm 2 difference) in TCA I1-M2 value, but more similar with Late 
Pleistocene North Europe (23 mm 2 difference, for available TCA 
I1-M3 data). Both size and shape components indicate Gough's 
Cave is dissimilar to Anglo-Saxon, and even less similar to 
Recent Britain. The DAS data, however, suggest close morpho- 
logical similarities between Gough's Cave and early North Europe. 
The discrepancies may reflect temporal fluctuations in environ- 
ment/subsistence, with the metric data more sensitive than 
morphology to these variables. In addition, sexual dimorphism 
and apportionment of tooth size may also have an effect. 

Pathology/occlusal attrition/crown chipping: Lack of carious 
teeth, periodontal pathology and low instance of enamel defects in 
Gough's Cave is well within the range of other hunter/gatherer 
populations (Cook & Buikstra 1979; Leigh 1925; Turner 1979). The 
one individual with less than 2 mm of root exposure between CEJ 
and the alveolar border is not indicative of periodontal disease, but is 
most likely the result of further root eruption to compensate for 
attrition, and correlated with age (Clarke & Hirsch 199 1 ). Similarly, 
there is one young adult with antemortem tooth loss, suggesting an 
occupationally related cause rather than due to carious activity or 
periodontal disease. 



32 



D.E. HAWKEY 



Table 8 Incisor Breadth ratio (IB) of upper central and lateral incisors compared with early and recent populations. 



IB 



Area/Site 



Sex 



Source 



0.83 

0.83 
0.80 
0.80 
0.80 
0.78 
0.78 
0.75 
0.72 



Gough's Cave (Late Paleolithic-Mesolithic) 

Jomon (Early Japan) 
Mehrgarh (Neolithic Pakistan) 
Mahadaha (Mesolithic India) 
Nubia (Mesolithic East Africa) 
Khoisan (Recent South Africa) 
Anglo-Saxon (Early Britain) 
Natufian (Epi-Paleolithic Levant) 
Britain (Recent) 



M&F 

M 
M&F 
M&F 

M 

M 

M 
M&F 

M 



Present study 

Brace & Nagai 1982 

Lukacs 1985 

Lukacs & Hemphill 1992 

Calcagno 1986 

Haeussler et al 1989; van Reenan 1982 

Lavelle 1968 

Dahlberg 1960 

Lavelle 1972 



Table 9 Molar Crown Area calculated for upper and lower M 1 and M2 (Ml -M2CA) and compared to early and recent populations. 



Ml-M2CA(mm 2 ) Area/Site 



Sex 



Source 



536 
503 
503 
486 
486 
485 
465 
448 
428 



Nubia (Mesolithic East Africa) 

Mahadaha (Mesolithic India) 

Natufian (Epi-Paleolithic Levant) 

Mehrgarh (Neolithic Pakistan) 

Gough's Cave (Late Paleolithic-Mesolithic) 

Britain (Recent) 

Jomon (Early Japan) 

Anglo-Saxon (Early Britain) 

Khoisan (Recent South Africa) 



M 
M&F 
M&F 
M&F 
M&F 

M 

M 

M 

M 



Calcagno 1986 

Lukacs & Hemphill 1992 

Dahlberg 1960 

Lukacs & Hemphill 1991 

Present study 

Lavelle 1968 

Brace & Nagai 1982 

Lavelle 1968 

Haeussler et al 1989; van Reenan 1982 



Table 10 Comparisons of Gough's Cave metric and morphological data forTCA absolute mean difference (TCAD), Penrose Size (PEN SIZE), Penrose 
Shape (PEN SHAPE), and Morphology (DAS) ranked in terms of most similar to least similar. 



Site/ Area 


TCAD 


Site/Area 


PEN SIZE 


Site/Area 


PEN SHAPE 


Site/ Area 


DAS% 


Mehrgarh 


3 


Natufian 


0.02 


Natufian 


0.87 


Early Europe 


83.3 


Natufian 


20 


Mehrgarh 


0.04 


Mehrgarh 


0.97 


Early S. Asia 


83.3 


Early Europe 


23 


Mahadaha 


0.05 


Jomon 


1.07 


Early Jomon 


80.0 


Jomon 


53 


Anglo-Saxon 


0.33 


Nubia 


1.10 


Recent Europe 


72.2 


Anglo-Saxon 


53 


Jomon 


0.39 


Mahadaha 


1.24 


Early Natufian 


70.0 


Recent Britain 


68 


Nubia 


0.59 


Khoisan 


1.34 


Early N. Africa 


70.0 


Mahadaha 


69 


Recent Britain 


0.97 


Anglo-Saxon 


2.39 


Recent S. Africa 


60.0 


Nubia 


124 


Khoisan 


1.14 


Recent Britain 


4.22 


Early Baikal 


52.6 


Khoisan 


124 










Early Nubia 


42.1 



Although an increase in the presence of calculus has been observed 
with the advent of Neolithic culture (Hildebolt & Molnar 1991), 
calculus deposits on the teeth are also found in populations with 
hunter-gatherer or mixed economies, and may actually be under- 
reported in archaeological specimens due to preservation or 
postmorten damage (Brothwell 198 1 ). Evidence ofphytoliths within 
the calculus deposits of Gough's Cave teeth has been recovered by K. 
Dobney of University of Bradford (reported in. Currant et al 1989). 
While the presence ofphytoliths can introduce a somewhat abrasive 
element into the diet, the teeth of the Gough's Cave sample are not 
excessively worn. However, a hunter-gatherer subsistence strategy 
also includes a reliance on meat, an element that is not necessarily 
abrasive to the dentition (Hillson 1986). Thus, the presence of crown 
microtrauma in Gough's Cave may be at least partially related to 
subsistence, especially when grit and bone may be present in the diet 
(Turner & Cadien 1969). 

Although caution should be used in a macroscopic analysis of 
enamel disturbances (Hillson & Brand 1997), it has been suggested 
that hunter/gatherers tend to be less severely affected by enamel 
hypoplasias (Cook & Buikstra 1979; Lukacs et al 1982), with the 
average age of onset between four to five years of age (Schulz & 
McHenry 1975). But the low instance of enamel hypoplasia in the 



Gough's Cave sample may not reflect a lack of nutritional stress, 
because there can be a variety of underlying causes (Goodman & 
Rose 1 99 1 ). The Mesolithic site of Mahadaha, for example, exhibits 
a high frequency (64%) of enamel defects, although the population 
appears to be free of nutritional stress markers in the osseous remains 
(Lovell 1992). Some authors have suggested that the amount of 
stress seen in a population may be reflected in greater dental asym- 
metry (Bailit et al 1970), or even by significant tooth size variation 
within age groups (Guargliardo 1982), neither of which are espe- 
cially apparent in the limited number of individuals from Gough's 
Cave. 

Other features: An absence of evidence for either enamel clean- 
ing striations or interproximal grooves, coupled with a lack of caries, 
periodontal pathology and only slight degree of calculus, suggest 
that the people of Gough's Cave did not need to practice a rigorous 
form of dental hygiene. Early teeth cleaning practices, however, have 
been noted in the Middle East, Asia, Africa, and North American 
Indians, who often utilized the frayed end of twigs to clean the teeth 
(Hawkey et al n.d.). Similarly there is even earlier evidence for 
interproximal grooves between the teeth, usually attributed to use of 
a 'toothpick' to remove irritating substances. These grooves have 
been reported for a variety of groups in Europe, dating from the Late 



HUMAN DENTAL REMAINS FROM GOUGH'S CAVE 



33 



Paleolithic to the Bronze Age (Alexandersen 1978; Bennike 1985; 
Formicola 1988; Frayer& Russell 1987;Turner 1988), were noted in 
the Neolithic remains from Mehrgarh (Lukacs & Pastor 1988), and 
possibly in South African Late Pleistocene sites (Grine & 
Henshilwood 2002; Grine et al 2000). 

The earliest evidence of intentional modification of the anterior 
teeth is the ablation seen at Minatogawa, dating to circa 1 8,000 years 
BP (Hanihara & Ueda 1982). In addition, intentional filing of the 
labial surface of incisors has been reported in early Holocene in 
South Asia (Kennedy et al 1981), and the practice of dental modifi- 
cation commonly occurs in Africa, the Americas, South Asia, Japan, 
Southeast Asia, Australia and Melanesia (Hawkey n.d b\ Milner & 
Larsen 1991 ). There are no instances of intentional dental modifica- 
tion (ablation, filing, or inlay) in the Gough's Cave sample, a fact that 
is supported by ethnographic reports that suggest the later popul- 
ations of Europe and the Middle East abhorred the loss of the anterior 
teeth (Guerini 1977; Kanner 1928). 

There are only a few cases of possible dental modification in early 
Britain (Jackson 1915), from two sites ascribed to a Neolithic culture 
(Dog Holes cave in Lancashire, and Perthi Chwareu caves in North 
Wales). Jackson's description of the specimens remains unconvinc- 
ing, however, as examples of intentional dental modification. There 
is an abnormal amount of wear on all four specimens, particularly 
those with loss of central incisors, and the loss may due to excessive 
attrition leading to exposure of the pulp chamber and premature 
exfoliation of the teeth. Interestingly, two of Jackson's specimens 
display antemortem loss of lower premolars; one individual from 
Gough's Cave has antemortem loss of LP 4 . Because the loss seen in 
both Gough's Cave and two of Jackson's specimens are not anterior 
teeth, it is unlikely to be intentional dental modification. The ante- 
mortem tooth loss seen in Gough's Cave, in particular, is more likely 
due to activity-induced traumatic injury, a situation observed in 
populations as ecologically disparate as the Arctic (Merbs 1983) and 
Pakistan (Lukacs & Hemphill 1990). 



CONCLUSIONS 

Although the dental remains from Gough's Cave are from a numeri- 
cally limited series, several trends are suggested, with the underlying 
assumption that dentition from these individuals accurately repres- 
ent the Late Pleistocene/Early Holocene populations of the British 
Isles. It is cautioned , however, that the results are tentative and may 
reflect statistical fluctuations due to small sample size. 

1 ) Morphology: The individuals from Gough's Cave have a sim- 
plified dental pattern, similar to the dentition of other Late 
Pleistocene/Early Holocene populations of North Europe, the 
Levant, and North Africa. They have similarities with two other 
groups, also with a simplified pattern: South Asia (Indodont 
pattern) and the Jomon (Sundadont pattern). They are dentally 
unlike populations of modern sub-Saharan Africa, Mesolithic 
Nubia, or the more complex Sinodont dentition of Lake Baikal. 
Gough's Cave lacks expression of any of the archaic traits, with 
the exception of P, Tomes' root. 

2) Metrics: Gough's Cave dentition is more similar in crown 
size to other Mesolithic European populations, exhibiting a 
significant reduction in tooth size from European 
Neanderthals, consistent with the post-Pleistocene trend in 
dental reduction. Among the Late Pleistocene/Early Holocene 
comparative samples, Gough's Cave is most similar in dental 
crown size to early populations from the Levant, South/South- 



west Asia, and North Europe, but unlike both early East Africa 
(Nubia) and modern sub-Saharan Africa (Khoisan). When both 
morphology and metric differences are compared, a similar 
pattern tends to occur, although no published metric data are 
available for North Africa (Iberomaurusian). There are tempo- 
ral differences within the British Isles: Gough's Cave appears 
most similar to the North Europe sample dating to the approxi- 
mately the same time period. Gough's Cave is less similar to 
Anglo-Saxon, with the Recent Britain sample even more dis- 
similar. A trend towards lateral incisor reduction occurs in 
later British populations, with Molar Crown Area remaining 
approximately the same as Gough's Cave. This finding may 
have some effect on the odontometric analysis, reflecting ap- 
portionment changes between the incisor/canine and premolar/ 
molar fields with time. 

3 ) Pathology/occlusal attrition/crown chipping: The dental path- 
ology profile is consistent with that of a hunter-gatherer lifeway, 
with absence of caries, no periodontal disease, and low fre- 
quency of enamel hypoplasia. The diet was probably not 
particularly abrasive, because the teeth show evidence of a 
gradual progression of attrition with age, rather than evidence of 
excessive wear during adolescence. An almost complete absence 
of enamel hypoplasia, along with little dental size asymmetry 
suggest a relatively healthy population. This low incidence of 
enamel hypoplasia may indicate a lack of nutritional stress, 
similar to that noted by Kennedy etal(\ 986), for the Mesolithic 
site of Sarai Nahar Rai in India, where hunter/gatherer subsist- 
ence strategy and ecological conditions may well have provided 
an abundance of food resources. Given the absence of caries in 
these remains, it is probable that the only instance of antemortem 
tooth loss in one individual may be occupationally related, 
especially considering the excessive enamel microtrauma found 
on the anterior teeth. 

4) Other features: Similar to other European populations, there 
is no convincing evidence of intentional dental modification. 
Although there have been some reports of interproximal 
'toothpick' grooves and cleaning striations among European 
Neanderthal populations, the lack of these features in Gough's 
Cave individuals may be related to the low instance of caries, 
and the presence of only slight-moderate degree of supra- 
gingival calculus. 



ACKNOWLEDGEMENTS: The author particularly wishes to thank 
Christopher B. Stringer and Louise T. Humphrey for their assistance (and 
patience). Special thanks are also due to Robert Kruszynski, Steven E. 
Churchill, and Christy G. Turner II. Dental morphology data for the South 
Asian samples were collected courtesy of funding from a National Science 
Foundation Dissertation Grant (# 9318334) and an American Institute of 
Indian Studies Junior Fellowship (1993-94). 



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Bull. nat. Hist. Mus. Lond. (Geol.) 58(supp): 7>1-AA 



Issued 26 June 2003 



Gough's Cave 1 (Somerset, England): an 
assessment of body size and shape 

TRENTON W. HOLLIDAY 

Department of Anthropology, Tulane University, New Orleans LA 70118, USA 

STEVEN E. CHURCHILL 

Department of Biological Anthropology and Anatomy, Duke University, Durham NC 27710, USA 

Synopsis. Stature, body mass, and body proportions are evaluated for the Cheddar Man (Gough's Cave 1 ) skeleton. Like many 
of his Mesolithic contemporaries, Gough's Cave 1 evinces relatively short estimated stature (ca. 166.2 cm [5' 5']) and low body 
mass (ca. 66 kg [ 1 46 lbs] ). In body shape, he is similar to recent Europeans for most proportional indices. He differs, however, from 
most recent Europeans in his high crural index and tibial length/trunk height indices. Thus, while Gough's Cave 1 is characterized 
by a total morphological pattern considered 'cold-adapted' , these latter two traits may be interpreted as evidence of a large African 
role in the origins of anatomically modern Europeans. 



INTRODUCTION 



Reconstructed stature, body mass, and body shape are all variables 
of interest in any attempt to understand the paleobiology of prehis- 
toric humans such as the 'Cheddar Man', or the Gough's Cave 1 
specimen. The relative completeness of the Gough's Cave 1 
postcranial skeleton allows each of these variables to be accurately 
reconstructed. Such variables are of interest both for evolutionary 
and non-evolutionary questions. For example, any body mass and/or 
stature differences between Mesolithic humans, such as Gough's 
Cave 1, and recent humans are unlikely to be evolutionary in nature. 
Nonetheless, they are of interest to paleobiologists since they may 
reflect the nutritional and overall health status of prehistoric popul- 
ations. In contrast, body proportions vary among recent humans, 
presumably as the result of climatic selection. Yet body proportions 
appear to have a large genetic component, and, over evolutionarily 
short periods, since they are the result of apparently long-term 
climatic selection (based on migrant studies), they may provide 
evidence of population movements or migration from different 
climatic regimes (Holliday, 1997a). 

Stature in Gough's Cave 1 is predicted from lower limb long bone 
lengths using Trotter and Gleser's (1958) standard formulae for 
Euroamericans (discussed in detail below). Body mass for the speci- 
men is predicted using two methods outlined in Ruff et al. (1997). 
The first method involves computing the arithmetic average of 
predictions based on three separate body mass/femoral head diameter 
regressions derived from recent human skeletal material. In the 
second method, body mass is predicted from stature and bi-iliac 
breadth. In concert these two variables (stature and bi-iliac breadth) 
are known to provide an accurate estimate of body mass in living 
humans, and have an added advantage in that they are independent of 
the locomotor biomechanical stresses to which the femoral head is 
subject (Ruff et al., 1997). Ruff's stature/bi-iliac breadth predictive 
formula is derived from data on living humans. Therefore, in order to 
use this method with fossils, stature was estimated using the Trotter 
and Gleser (1958) formulae, and a 5% correction factor was added to 
bi-iliac breadth to account for soft tissue. All formulae used to 
predict body mass were kindly provided by Prof. C.B. Ruff. 

With regard to body shape or proportions, there are several means 
by which these features may be accurately reconstructed from 
skeletal remains; these means approximate some of the anthro- 



pometric data taken on living human subjects. The measures that are 
used in this study reflect the following: 1 ) intralimb proportions (i.e., 
relative lengths of the proximal and distal limb segments), 2) limb/ 
trunk proportions, 3) body linearity relative to overall body mass, 
and 4) body breadth relative to stature. For all analyses, Gough's 
Cave 1 is compared to other Late Pleistocene and Early Holocene 
associated skeletons as well as to a large sample of recent humans 
from across the western Old World (Africa and Europe). The fossils 
have been placed into Mesolithic (< 10,000 BP), Late Upper 
Paleolithic (LUP; 1 1,000-19,000 BP), Early Upper Paleolithic (EUP; 
20,000-28,000 BP) and Neandertal (> 30,000 BP) samples, while 
the recent humans have been placed into three geographical 
subsamples: Europe, North Africa and Sub-Saharan Africa. Detailed 
discussion of these samples is found in Holliday (1995). 



BODY SIZE 

Stature 

For all samples, stature was predicted using Trotter and Gleser's 
( 1 958 ) formulae for the tibia and femur; if both bones were present, the 
mean of the two resultant predictions was used. Formulae for Euro- 
american males were used for Gough's Cave 1 and all comparative 
samples, with the exception of the recent Sub-Saharan Africans and 
the European EUP, for whom African- American formulae were used. 
These regression formulae are more appropriate because these two 
groups have a demonstrably more 'tropically-adapted' body shape 
which is more similar to that of African-Americans (Holliday, 1997a). 
Table 1 presents summary statistics for predicted stature among 
Gough's Cave 1 and the comparative samples. The Gough's Cave 
specimen has a predicted stature of 166.2 cm, which falls just below 
the Mesolithic male mean of 1 67.5 cm. His predicted stature is much 
shorter relative to recent European males; he falls below their 25th 
percentile. Importantly, predicted stature values for the fossils are 
similar to those given in Frayer (1984), who used many of the same 
specimens, but temporally subdivided his samples differently than 
has been done here. As an example of the similarity of results, 
Frayer's (1984) Mesolithic sample had a predicted male stature of 
167.8 cm, almost identical to our mean of 1 67.5 cm. Also, his Upper 
Paleolithic male mean of 174.3 cm is somewhat (although not 



© The Natural History Museum, 2003 



38 



T.W. HOLLIDAY AND S.E. CHURCHILL 



Table 1 Summary statistics (mean, standard devition, number of 

specimens) for Gough's Cave 1, fossil and recent human male samples - 
predicted stature (in cm). 

Predicted stature 



Gough's Cave 1 

European Mesolithic 

European Late Upper Palaeolithic 

European Early Upper Palaeolithic 

European Neandertals 

Recent Europeans 

Recent North Africans 

Recent Sub-Saharans 



166.2 

167.5,4.8,7 

170.2,6.6,17 

170.1,7.9,11 

166.7,3.8,4 

171.6,5.8,311 

167.4, 5.9, 75 

164.7,8.2,62 



statistically significantly) higher than our LUP and EUP means of 
170.2 cm and 170.1 cm, respectively. 

Not unexpectedly, our results suggest that stature in Europe is 
highest among Upper Paleolithic (both EUP and LUP) and recent 
Europeans. Neandertals and Mesolithic Europeans, on the other 
hand, are significantly shorter than recent Europeans (two-tailed t 
test, p = 0.020 and 0.048, respectively). These results are similar to 
those reported by Frayer et al. (1993) and Formicola & Giannecchini 
(1999). With regard to the Mesolithic sample, this reduction in 
stature may be due to a drop in dietary protein. Such a drop could 
have followed decreased reliance on big game following the refor- 
estation of Europe, a phenomenon documented by archaeologists for 
many early Holocene hunter-gatherers (Straus et al., 1980; Geddes 
etal., 1986). 

Note that among the recent human groups, stature appears to 
decrease as one moves toward the equator. This is likely a secondary 
consequence of a decrease in body mass associated with increas- 
ingly hotter, more tropical temperatures, following Bergmann's rule 
(see below). 

Body Mass 

Table 2 gives predicted body mass summary statistics for Gough's 
Cave 1 and the comparative male sample. Among the recent human 
samples, there is a clear decrease in body mass (based on either 
predictive method) from higher to lower latitudes. This reflects 
adherence of humans to Bergmann's (1847) ecological rule (dis- 
cussed below). The Gough's Cave 1 specimen has a predicted body 
mass of 64.8 kg based on femoral head size, and a mass of 67.3 kg 
based on stature and bi-iliac breadth. It is noteworthy that despite the 
fact that the two methods use very different anatomical features, the 
two predictions deviate from each other by less than 4%. Note, also 
that across all groups, the mean body mass estimates using the non- 
biomechanical (stature/bi-iliac breadth) method are close to those 
derived from the femoral head. The greatest difference between the 
two methods is found among the EUP sample, whose body mass 



prediction based on stature and bi-iliac breadth is 5.8% higher than 
the one based on femoral head diameter. Note, too, that while the 
Neandertal sample appears to be characterized by high body mass, 
there is relatively little evidence for a subsequent change in body 
mass in Europe from the EUP to the present (a result consistent with 
the findings of Ruff et al., 1997). As for the specimen of interest, 
Gough's Cave 1 is not atypical among early Holocene Europeans in 
mass; he falls slightly below the Mesolithic male mean based on the 
femoral head prediction, and slightly above the mean for the stature/ 
bi-iliac breadth prediction. He, like most of his Mesolithic cohorts, is 
light relative to recent Europeans; his femoral head-predicted and bi- 
iliac breadth/femoral length predicted weights fall on the 29 lh and 
37 th recent European male percentiles, respectively. 



BODY SHAPE 

Intralimb Proportions 

Elongation of the distal limb segment relative to the proximal has 
been demonstrated to be associated with overall limb elongation in 
both the upper and lower limb (Meadows & Jantz, 1995), and is 
correlated with climatic variables (Roberts, 1978; Trinkaus, 1981). 
Distal limb segment elongation is typically quantified in the form of 
brachial (radius length/humeral length x 100) and crural (tibial 
length/femoral length x 100) indices. These skeletal measures are 
comparable to the anthropometric antebrachial index (forearm length/ 
upper arm length x 100) and calf/thigh index (calf length/thigh 
length x 100), respectively, which are commonly taken on living 
people (Roberts, 1978). 

Table 3 gives summary statistics for the brachial and crural indices 
of the Gough's Cave 1 specimen and fossil and recent human 
samples. Note that among the recent humans, the indices show a 
cline from lower to higher latitudes, with high indices in the former, 
and low indices in the latter. This is presumably the result of long- 
term climatic selection (discussed below). Within groups, male and 
female brachial and crural index values are similar (males do, 
however, tend to have higher brachial indices than females; Trinkaus, 
1981 ; Holliday, 1995). Given the difficulty in assigning sex to some 
fossil specimens (as well as the already small size of the fossil 
sample), combined-sex means are given in Table 3. This does not 
affect the overall pattern, as will be evident below, when we discuss 
Gough Cave l's relationship to other males from the comparative 
sample. 

As is evident from Table 3, the Cheddar specimen, like other Late 
Pleistocene and early Holocene Europeans, has elongated distal limb 
segments in both the upper and lower limb. In fact, Gough's Cave 1 
has indices not unlike the means of the recent African samples, and 



Table 2 Summary statistics for Gough's Cave 1, fossil and recent human 
males - predicted body mass (in kg). 



Table 3 Summary statistics for Gough's Cave 1 , fossil and recent human 
samples - brachial and crural indices. 





Femoral Head Method 


Stature/BIB Method 




Brachial Index 


Crural Index 


Gough's Cave 1 


64.8 


67.3 


Gough's Cave 1 


77.1 


88.9 


European Mesolithic 


66.9, 7.2, 7 


66.0, 2.3, 6 


European Mesolithic 


77.5, 1.9, 10 


85.5,2.6, 10 


European Late Upper 


67.7, 6.6, 14 


67.4, 8.2, 6 


European Late Upper 


78.6,3.0, 17 


85.1,1.9,22 


Palaeolithic 






Palaeolithic 






European Early Upper 


65.8, 10.0, 10 


69.6, 7.3, 6 


European Early Upper 


77.9,2.2, 17 


85.4, 1.9, 13 


Palaeolithic 






Palaeolithic 






European Neandertals 


82.9, 4.3, 4 


79.3,-1 


European Neandertals 


73.2,2.5,5 


78.7, 1.6,4 


Recent Europeans 


69.3,7.3,134 


71.0,7.4, 126 


Recent Europeans 


75.0,2.5,391 


82.7, 2.4, 436 


Recent North Africans 


59.0, 7.6, 73 


61.3,5.5,60 


Recent North Africans 


78.6,2.4, 136 


85.0,2.3,133 


Recent Sub-Saharans 


54.7, 8.5, 53 


53.6,8.6,49 


Recent Sub-Saharans 


78.6,2.8, 103 


85.4,2.4,110 



GOUGH'S CAVE 1 : ASSESSMENT OF BODY SIZE AND SHAPE 



39 



his crural index value actually falls above the Sub-Saharan African 
mean. It is not, however, too unusual to find a male European 
specimen today with a brachial index value equal to or higher than 
that of the Gough's Cave specimen; Gough's Cave 1 falls right on the 
75th percentile for the recent European males (n = 239). However, 
his crural index would be extremely unusual in a sample of recent 
Europeans, since his value falls above the 99th percentile for recent 
European males (n = 273). 

His values are not, however, unusual among European Mesolithic 
(nor Paleolithic) humans. His brachial index is virtually identical to 
the Mesolithic mean, and while his crural index is above the 
Mesolithic mean, one of the 10 Mesolithic specimens sampled 
(Teviec 11) has yet a higher crural index (89.1). 

Limb/Trunk Proportions 

The fact that limb/trunk proportions of modern humans covary with 
climate and geography has been documented for both skeletal ( Holliday, 
1995, 1997a) and anthropometric samples (via the relative sitting 
height index { sitting height/stature x 100}, Roberts, 1978). Given the 
largely complete (albeit poorly reconstructed) vertebral column of the 
Gough's Cave specimen, one can estimate skeletal trunk height ( STH 
= summed dorsal vertebral elements T1-L5 + sacral ventral length; 
Franciscus & Holliday, 1992) as a body size or trunk length variable 
to which relative limb length may be assessed. As was done for the 
thoracic and lumbar vertebral column heights (Chuchill & Holliday, 
2002), STH is estimated from those vertebral elements preserved in 
Gough's Cave 1 , using a least-squares regression foracomplete recent 
human series (n = 45). The formula used is: Y = 1 .086.V - 1 .806; r 2 = 
0.998, where x (partial trunk height, or PTH) is the summed dorsal 
body heights for T4-L5, sacral ventral length, and the ventral body 
height of Tl . The 'reconstruction' for display of the specimen neces- 
sitated further estimation. Thoracic vertebrae 6 and 7 were glued 
together with a mock intervertebral disk between them; thus their 
combined dorsal height was measured and the height of the interven- 
ing 'disk' (2.9 mm) was subtracted, yielding 39.8 as the estimate of 
combined T6-T7 dorsal height. The combined height of T8-T9 (42.5) 
andTl 1-T12 (48.1 ) were estimated in the same manner. The predic- 
tive equation based on the above measurements yields an STH of 
483.9 mm, with a SE of the estimate of 1 .6 mm. The 95% confidence 
limits for the prediction are 480.6-487.2 mm, a span which is only 
1.4% of the prediction itself, indicating that STH can be accurately 
predicted in Gough's Cave 1. 

As discussed in Churchill & Holliday (2002), the height of Ched- 
dar Man's vertebral column (as reflected in thoracic and lumbar 
column heights) was short for a Mesolithic male. Thus, it is not 
surprising that the Gough's Cave specimen possesses a short STH, as 
well. The specimen's STH of 483.9 falls well below (although within 
one standard deviation of) the Mesolithic male mean of 5 1 1 .6 (n = 4), 
and only one Mesolithic male, the diminutive Hoedic 9, has a shorter 
trunk. However, the most important question that remains is how 
Gough's Cave 1 compares in terms of limb length relative to trunk 
height. In order to elucidate these patterns, limb segment length 
(maximum humeral, radius and tibial length and femoral bicondylar 
length) to trunk height ratios were computed for the comparative 
fossil and recent human sample, and are compared to Gough's Cave 
1 in Tables 4 and 5. Sexual dimorphism in these traits exists, but is 
minimal (Holliday, 1995); thus, as was done with the brachial and 
crural indices, Gough's Cave 1 is compared to combined-sex sam- 
ples. As was evident in intralimb proportions, among recent humans 
there is a clinal distribution of limb/trunk ratios, with Sub-Saharan 
Africans exhibiting the highest mean indices, the Europeans the 
lowest, and North Africans intermediate between the two groups. 



Table 4 Summary statistics for Gough's Cave 1 , fossil and recent human 
samples - upper limb segment/trunk height ratios. 



HL/STH 



RL/STH 



Gough's Cave 1 66.7 51.5 

European Mesolithic 61.7,3.7,7 47.9,2.7,7 

European Late Upper Palaeolithic 61.2, 2.8, 15 48.3, 2.4, 12 

European Early Upper Palaeolithic 69.1,4.0,8 55.0,2.7,7 

European Neandertals 64.0, 1.5,3 47.0, 0.2, 3 

Recent Europeans 63.6, 3.4, 124 47.9, 2.8, 123 

Recent North Africans 66.0, 3.8, 62 51 .9, 3.4, 62 

Recent Sub-Saharans 69.6,4.1,51 55.0,4.0,51 



Table 5 Summary statistics for Gough's Cave 1, fossil and recent human 
samples - lower limb segment/trunk height ratios. 



FL/STH 



TL/STH 



Gough's Cave 1 89.7 79.8 

European Mesolithic 87.4, 3.9, 7 74.0, 4.0, 7 

European Late Upper Palaeolithic 86.6, 3.4, 15 73.6, 3.5, 13 

European Early Upper Palaeolithic 96.0, 5.1,7 84.0, 4.6, 6 

European Neandertals 89. 1 , 0.0, 2 7 1 .2, 1 .0, 2 

Recent Europeans 88.6,4.4,123 73.6,4.3,124 

Recent North Africans 94.2, 5.5, 63 79.8, 4.9, 60 

Recent Sub-Saharans 97.7,7.5,51 84.1,6.5,51 

For the upper limb/trunk height ratios (Table 4), Gough's Cave 1 
differs not only from recent Europeans, but from Late Upper 
Paleolithic (LUP) Europeans, as well. In fact, relative to trunk 
height, the Cheddar specimen is somewhat long-armed, and is most 
similar to the recent North Africans in this regard. He is less long- 
armed, however, than the average recent Sub-Saharan African or 
European Early Upper Paleolithic (EUP) humans. While the distri- 
bution of sample means provides an overall pattern of differences, 
we may still ask how unusual would upper limb/trunk height ratios 
equal to or greater than that of Gough's Cave 1 be among recent 
Europeans? An examination of the male European distribution pro- 
vides some insight. For the humeral length/trunk height ratio, he falls 
on the 75% percentile of recent European males, while for the radius 
length/trunk height ratio, he falls above the 85% percentile. Thus 
while he does exhibit a positive deviation from the mean, sampling 
a recent European male who shares his upper limb/trunk height (or 
greater) values could be as common as 1 in 4. 

The lower limb/trunk height ratios reveal a slightly different 
pattern (Table 5). The Cheddar specimen's femoral length/trunk 
height ratio is very similar to the recent European mean, while his 
tibial length/trunk height ratio is 2.5 standard deviations above the 
recent European mean - indicating that he has an extremely long 
tibia relative to the height of his trunk. His percentile placement 
among the recent Europeans males reflects this dichotomy; he falls 
on their 60th percentile for the femoral length index, and above the 
94th percentile for the tibial length index. With regard to the recent 
Africans, he falls below the Sub-Saharan African mean for both 
indices, and below the North African FL/STH mean. His TL/STH 
value, however, is identical to the North African average. 

In comparison with other European fossils, Gough's Cave 1 
possesses a relatively longer femur than the mean of all but one fossil 
sample (the long-limbed EUP), although he falls well within the 
range of all but the short-limbed Neandertal samples. His high 
relative tibial length index, however, is somewhat more unusual in 
the sense that he exceeds the range of the LUP sample, and, addition- 
ally, he evinces the highest TL/STH index of the Mesolithic sample. 
In fact, with regard to relative tibial length, among the fossil groups 
only the long-limbed EUP sample exceeds his value. 



40 



T.W. HOLLIDAY AND S.E. CHURCHILL 



Body Linearity Relative to Mass 

Another body shape feature known to covary with climate is relative 
body linearity. In living populations, the weight: height, orponderal, 
index is used as a measure of this relationship (e.g., Newman, 1961 ; 
Schreider, 1964, 1975; Eveleth, 1966; Hiernaux etal, 1975). This 
relationship is most easily quantified skeletally via relative femoral 
head size (i.e., antero-posterior femoral head diameter/femoral 
bicondylar length x 1 00). This index should reflect relative linearity, 
since the femoral head is highly correlated with body mass, while 
femoral length is highly correlated with stature. This skeletal index 
was (not surprisingly) found to vary significantly between males and 
females, with males possessing relatively larger femoral heads than 
females (two-tailed / test, p < 0.0001 ), and thus Gough's Cave 1 is 
compared only to other males for this trait. 

Table 6 reports the summary statistics for this trait among the 
comparative samples and the Cheddar specimen. Within the recent 
humans, there is a clear clinal pattern from Sub-Saharan Africa 
through North Africa and into Europe, such that the femoral head 
becomes relatively larger with increasing latitude (see also Ruff, 
1994). With regard to fossil humans, note the extremely high indices 
exhibited by the male Neandertals. For this index, both Neandertal 
males (La Chapelle-aux-Saints 1 and La Ferrassie 1 ) fall beyond the 
99th percentile of recent European males (n = 134). The other 
European fossils, including the Mesolithic males and the Gough's 
Cave 1 specimen himself are virtually identical to recent Europeans 
for this trait. Only the EUP sample slightly deviates from the 
European pattern of relatively large femoral heads; they are more 
similar to recent North Africans in that their femoral heads are 
somewhat smaller (although not as small as those of the Sub-Saharan 
Africans). 

Body Breadth Relative to Stature 

Bi-iliac breadth, or bi-cristal breadth, as it is sometimes called, is 
measured as the transverse diameter of the superior margin of the 
pelvic girdle. This raw measurement is correlated with climatic 
variables (Crognier, 1981 ; Ruff, 1994), but its fit with climate and/or 
geography significantly improves when it is scaled to a linear dimen- 
sion of the body such as stature (Roberts. 1978; Ruff, 1991. 1993, 
1994). For the samples presented here, stature is unknown, and 
therefore must be predicted from long bone length, e.g. femoral 
length. In such cases, then, predicted stature is each individual's 
femoral length subsequent to an arithmetic manipulation, (i.e., femo- 
ral length x slope, +Y-intercept). Such prediction formulae inevitably 
introduce error into the analysis, however, since biologically speak- 
ing, many individuals are expected to fall well above or well below 
the predictive line. Thus, to avoid the introduction of further error, 
stature is not predicted for this analysis, but rather, femoral length 
(which is highly correlated with stature) is used in its stead. 

The first means by which the body breadth to height relationship 
can be investigated is via the computation of ratios - in this case, bi- 
iliac breadth / femoral bicondylar length x 100. Due to the fact that 
females have wider trunks relative to stature than do males, the 
values for this index are significantly different between the sexes 
(two-tailed t test, p < 0.000 1 ), and therefore the Cheddar specimen is 
compared solely to males for this variable. Table 6 reports the 
summary statistics for the males in the comparative sample and the 
Cheddar specimen. Gough's Cave 1 lies well within 1 standard 
deviation of the Mesolithic, LUP and recent European male means. 
Likewise, his value is only 1 .4 standard deviations above the North 
African mean. However, he falls over 3 standard deviations above the 
recent Sub-Saharan African mean; as discussed below, this group is 
characterized by some of the longest limbs and narrowest trunks of 



Table 6 Summary statistics for Gough's Cave 1 , fossil and recent human 
males - femoral head/femoral length ratios (FHAP/FL) and bi-iliac 
breadth/femoral length ratios (BIB/FL). 



FHAP/FL 



BIB/FL 



Gough's Cave 1 10.7 63.3 

European Mesolithic 10.7,0.5,6 62.1,3.2,6 

European Late Upper Palaeolithic 10.8,0.7,15 61.2.5.1.10 

European Early Upper Palaeolithic 10.1, 0.4, 1 56.6, 3.2, 6 

European Neandertals 12.3,0.4,4 69.8.-, 1 

Recent Europeans 10.6,0.5.134 61.2.3.4.126 

Recent North Africans 9.9,0.6,72 57.3,4.4,60 

Recent Sub-Saharans 9.5, 0.6, 53 52.6. 3.0, 49 

any humans. Interestingly, while based on extremely small samples, 
the earlier European fossil samples stand in marked contrast to each 
other and to recent Europeans. The Neandertals (albeit solely repre- 
sented by the La Chapelle-aux-Saints 1 specimen) are characterized 
by an extremely high index, indicative of their broad body breadth 
relative to stature (Ruff, 1991, 1993, 1994;Trinkause/a/., 1994).By 
way of contrast, the earliest modern European males (represented by 
6 individuals) have low indices; in fact, their mean index falls 
between those of the North and Sub-Saharan Africans. 

A second means of evaluating relative body breadth has been used 
extensively by Ruff (1991, 1993, 1994), and involves plotting rela- 
tive bi-iliac breadth indices, like those calculated above, against 
stature in bivariate space. Using this method, one can evaluate the 
relationship between the 'size-corrected' index and a measure of 
overall size (in Ruff's case, stature; here again, femoral length is 
used in its stead). Ruff has shown that among recent humans, there is 
little overlap among broad geographically circumscribed samples 
for this bivariate relationship, and thus this method could provide 
some insight into the relative position of the Cheddar specimen. 
Figure 1 is a scatter plot of the bi-iliac breadth/femoral length ratios 
regressed on femoral length for the recent Sub-Saharan Africans 
(squares), the recent Europeans (crosses) and Gough's Cave 1 (star). 
The lines fitted to the recent samples are least-squares regression 




350 



417 483 

Femoral Length 



Fig. 1 Scatter plot of bi-iliac breadth/femoral length index on femoral 
length. Recent Europeans are indicated by crosses; recent Sub-Saharan 
Africans by squares. Gough's Cave 1 is indicated by a star. The lines for 
the two recent human samples are least-squares regression lines. 



GOUGH'S CAVE 1 : ASSESSMENT OF BODY SIZE AND SHAPE 



41 



lines. As Ruff has found, there is good separation of the Europeans 
from the Sub-Saharan Africans throughout most of the size range. 
Informatively, Gough's Cave 1 falls squarely on the European re- 
gression line, far above the Sub-Saharan African line. 

Multivariate Assessment of Body Shape 

Any assessment of an individual's body size and proportions is at its 
base an assessment of that individual's total morphological pattern. 
While the individual analyses presented above when considered as a 
whole provide tantalizing clues as to the total morphological pattern 
of the Cheddar Man, these analyses are likely not as informative as 
would be a multivariate assessment based on the same morphologi- 
cal variables. In fact, a multivariate analysis may be expected to 
resolve some of the conflicting results obtained above. For example, 
in relative body linearity, relative body breadth and limb/trunk 
(excepting the tibia) proportions, the Gough's Cave specimen looks 
essentially like a recent European (albeit occasionally at the more 
linear end of the European range). In contrast, his tibia/trunk, bra- 
chial, and especially his crural index are more similar to those of 
more tropically-adapted groups (e.g., Africans). 

What then, is the total morphological pattern of body size and 
shape exhibited by Gough's Cave 1? The way to discover this is to 
investigate overall body proportions in multivariate space, taking the 
variances and covariances of all the skeletal manifestations of body 
shape into account. Once this is done, Gough's Cave 1 will either 
continue to fall among recent Europeans, or he could possibly 
exhibit a somewhat different, more tropically-adapted pattern. 

The variables to be used in the multivariate analysis and their 
abbreviations are found in Table 7. Note that these measurements are 
the same variables used to compute ratios and/or which were plotted 
in bivariate space. They should therefore provide an accurate reflec- 
tion of total body shape. The method chosen for body shape extraction 
is that outlined by Mosimann and colleagues (Mosimann & James, 
1979; Darroch & Mosimann, 1985; James & McCulloch, 1990). 
These morphometricians argue that an individual's overall size is 
best represented by the geometric mean of all the measurements 
taken on that individual. The geometric mean (or 'log size' as the 
authors denominate it) can then be used to create scale-free ratios, or 
'shape' variables, between each of the individual's measurements 
and his geometric mean. The utility of the shape variables lies not in 
the 'removal' of size per se, but in the ability of the researcher to 
determine if there is a relationship between size and shape via 
correlation analyses. The application of this method to anthropologi- 
cal data sets is discussed in greater detail elsewhere (e.g., Falsetti et 
ai, 1993; Jungers et ah, 1995). In this study, since the primary 
interest is the body shape of Cheddar Man, discussion is limited to 
the analysis of shape variables. The variance-covariance matrix 
(VCM) of the shape variables for a combined sample of fossil and 

Table 7 First two principal components of shape variables - fossil and 
recent humans. 

Eigenvector Coefficient 
I II 



Femoral A-P head diameter (FHAP) 
Bi-iliac breadth (BIB) 
Femoral bicondylar length (FL) 
Humeral maximum length (HL) 
Tibial maximum length (TL) 
Radius maximum length (RL) 
Skeletal trunk height (STH) 

Eigenvalue 

% total variance 



0.305 


-0.860 


0.591 


0.451 


-0.246 


0.070 


-0.178 


0.037 


-0.404 


0.124 


-0.421 


-0.009 


0.591 


0.187 


0.0094 


0.0032 


58.25 


19.63 






<u 

a 
a 



u 




PCI Shape (58.3%) 

Fig. 2 Scatter plot of PC2 on PC 1 (shape data). Crosses are recent 
Europeans, open squares are recent Sub-Saharan Africans, triangles are 
Early Upper Paleolithic, circles are Late Upper Paleolithic, closed 
squares are Mesolithic, star is Gough's Cave 1. Lines indicate range of 
the recent human samples. 



recent humans (n = 225), all of whom preserve the measurements in 
question, was computed and then subjected to principal components 
analysis (PCA). 

The eigenvector coefficients and eigenvalues for the first two 
principal components of the log shape data are found in Table 7. The 
first principal component (PCI) accounts for 58.3% of the total 
shape variance, and contrasts limb segment length (particularly the 
distal segments) with femoral head diameter, bi-iliac breadth and 
skeletal trunk height. The PC scores along this axis are not signifi- 
cantly correlated with overall size (i.e., the geometric mean; r 2 = 
0.008, p = 0. 1 758). PC 1 is readily interpreted as a climatic adaptation 
component, since it separates heavier, less linear individuals (more 
cold-adapted) from lighter, more linear individuals (more heat- 
adapted). The second principal component (PC2) accounts for 19.6% 
of the shape variance and contrasts bi-iliac breadth and trunk height 
with femoral head diameter. The scores along this axis are correlated 
with log size (r 2 = 0.18, p < 0.0001), and this component tends to 
segregate males (who on average have large femoral heads and 
relatively narrow pelves) from the small femoral-head possessing 
and wider hip bearing females (albeit with considerable overlap). 

Component scores for the European early modern fossils (includ- 
ing Gough's Cave 1 ), as well as the recent Sub-Saharan Africans and 
recent Europeans are plotted in Figure 2. Note that the separation of 
the groups is along the first principal axis. This axis contrasts 
individuals on the left, who possess short distal limbs, wide and 
relatively long trunks, and large femoral heads from those individu- 
als on the right, who are characterized by relatively short and narrow 
trunks, long distal limb segments and smaller femoral heads. There 
is no separation of the groups (fossil or recent) along the second 
principal axis. Note that for the first principal component, there is 
relatively little overlap between the recent Sub-Saharan Africans 
(represented by open squares ) and the recent Europeans (represented 
by crosses). Gough's Cave 1 (the star) and his contemporaries, the 
European Mesolithic specimens (indicated by dark squares) fall 
clearly among the recent Europeans, as do the LUP specimens 



42 



T.W. HOLLIDAY AND S.E. CHURCHILL 



(indicated by circles). Only 2 of the 9 (22%) LUP specimens (Barma 
Grande 2 and Bichon 1 ) even fall in the region of overlap between the 
recent Sub-Saharan Africans and Europeans, and none of the 
Mesolithic sample does. In contrast, there is a tendency for the EUP 
specimens (indicated by triangles) to fall among the Sub-Saharan 
Africans and outside of the European sample range. This specific 
result is said to be indicative of a relatively recent African origin for 
the earliest modern Europeans, and is discussed in detail elsewhere 
(Holliday, 1995, 1997a). What is of most interest to this chapter is 
that the Gough's Cave specimen, despite possessing some 'non- 
typically' European traits, falls squarely among the Europeans in 
multivariate space, albeit toward the more linearly-built end of the 
distribution. 

Discussion 

Gough's Cave 1 is relatively unremarkable with regard to stature and 
body mass; he is small, yet similar to all European samples, save the 
heavier Neandertals. However, his body shape poses some interest- 
ing contrasts which need to be further explored. Among recent 
humans, clear differences in body shape manifest themselves among 
geographically-dispersed samples. In terms of relative sitting height, 
for example, some Australian Aboriginal and Sub-Saharan African 
groups evince mean relative sitting height indices as low as 47.0, 
while at the other extreme, many Inuit (Eskimo) samples evince 
mean indices of around 54.0 (Eveleth & Tanner, 1976). What this 
means is that among some of the more tropically-adapted groups 
worldwide, the head, neck and trunk comprise less than half (ca. 47% 
or less) of a person's stature. Yet another way of looking at this is that 
among these groups, the lower limb accounts for more than half (ca. 
53+%) of a person's standing height. By way of contrast, among the 
Inuit and other cold-adapted groups, the head, neck and trunk make 
up well over half (ca. 54+%) of the average person's height, while the 
lower limbs make up considerably less than half (ca. 46% or less). 

The explanation for empirical patterns such as the above is that 
they are due to climatic selection, and more specifically, reflect the 
adherence of recent humans to the ecological 'rules' of Bergmann 
(1847) and Allen (1877). These rules state that within a widespread 
species of warm-blooded animals, those in colder regions will tend 
to be heavier (Bergmann's rule) and evince shorter extremities 
(Allen's rule) than do their more tropical conspecifics. Theoretically, 
it is argued that we find this pattern because animals in cold regions 
minimize their surface area: volume ratio (SA:V) in order to better 
conserve body heat, since heat loss occurs through the skin (i.e., the 
animal's surface). On the other hand, heat loss in hot environments 
may be facilitated by increasing relative surface area. Changes in 
body size and shape can drastically affect the SA:V ratio, as dis- 
cussed in Ruff (1994) and Holliday (1995). 

But do these rules apply to fossil humans as well, or is this an over- 
extention of biological uniformitarianism? Limited fossil data suggest 
that prehistoric human populations were characterized by 
ecogeographical clines that were perhaps even steeper than those 
one finds today (Trinkaus, 1981, 1991; Stringer, 1989; Ruff, 1991, 
1993, 1994). For example, the Kenyan Nariokotome Homo erectus 
skeleton (KNM-WT 15000) is said to be characterized by 'hyper- 
African' body proportions (Ruff and Walker, 1993), while European 
Neandertals are characterized by an extremely cold-adapted mor- 
phology (Trinkaus, 1981, 1986; Holliday, 1995, 1997Z>; Churchill, 
1998). 

How do the Gough's Cave 1 specimen and his contemporaries fit 
into this apparently climatically-driven geographical patterning? In 
order to address this question adequately, we must have at least some 
understanding of what the pattern in Europe was before the early 



Holocene, i.e., what was the temporal pattern of body proportions in 
the European Pleistocene? In other words, were there temporal trends 
in body shape during this time period? The answer is an emphatic 
'yes'. There is actually more temporal variability in body shape in 
Pleistocene Europe than there is spatial variability in the world today. 

We begin with the European Neandertals. They exhibit a clearly 
cold-adapted physique, including low brachial and crural indices, low 
limb/trunk ratios, extremely large femoral heads and wide trunks. 
Those who succeed them in the region, however, hominins differen- 
tially referred to as the 'Cro-Magnons' or the Early Upper Paleolithic 
(EUP) humans, exhibit the opposite pattern - high brachial and crural 
indices, high limb/trunk proportions, relatively smaller femoral heads 
and narrower trunks. Succeeding the Cro-Magnons are the Late Upper 
Paleolithic (LUP), and subsequent Mesolithic populations. These 
later two samples have, in this analysis, been divided at the Pleistocene/ 
Holocene boundary, with the Mesolithic sample (including Gough's 
Cave 1 ) being restricted to the latter epoch. This division may be 
biologically insignificant, however, since for virtually all analyses - 
univariate, bivariate or multivariate, the LUP and Mesolithic samples 
more closely resemble each other than they do any other group, fossil 
or recent (see also Holliday, 1995, 1997a). 

Combined or separate, the real question of interest is what was the 
pattern of body shape in LUP and Mesolithic humans? Importantly, 
the 'shared' morphology of these two samples (including the speci- 
men of interest) is in some regards paradoxical (Holliday, 1999). 
Late Upper Paleolithic and Mesolithic specimens retain the high 
brachial and crural indices of their presumed ancestors, the 'Cro- 
Magnons'. Yet unlike the Cro-Magnons, they tend not to possess 
relatively narrow trunks, relatively small femoral heads, or high 
limb-trunk ratios. Gough's Cave 1, as a general rule, follows this 
pattern. Like his contemporaries, his brachial and crural indices are 
near the upper extreme of the recent European sample. Likewise, as 
with others from his time period, his limb/trunk proportions are 
within the European range, although his values for HL/STH, RL/ 
STH and particularly his TL/STH indices are somewhat more ex- 
treme than those of average Europeans today. Recall that he falls on 
the 75th, 85th and 94th percentiles, respectively, of the recent 
European male sample for these traits. For the other traits (relative 
femoral head size and relative body breadth), however, he falls very 
near the recent European mean, and is distinctly different from 
recent Africans. In multivariate space, however, he lies within the 
European scatter, and beyond the range of recent Sub-Saharan 
Africans, as do his Mesolithic contemporaries and the majority of 
the LUP sample. By way of contrast, the EUP sample tends to cluster 
more closely with the recent Africans. 

Both in scientific articles (Frayer, 1992; Frayer et al., 1993) and 
the popular press (Shreeve. 1995), it has been pointed out that the 
retention of high brachial and crural indices among Late Upper 
Paleolithic and Mesolithic humans is problematic for Trinkaus' 
(1981) argument that these indices reflect elevated gene flow (or 
population dispersal) from Africa associated with the origins of 
modern humans. After all, these workers argue, the glacial cold of 
Europe should have modified, at least by the end of the Pleistocene, 
any previously incoming population toward a more cold-adapted 
morphology. Yet with regard to brachial and crural indices, the LUP 
sample have an even more extreme (almost 'hyper-tropical') mor- 
phology than their EUP forebears. 

As pointed out in Holliday (1999), this argument shows the 
problems that can arise when single traits are studied in isolation 1 . In 



'We can, for the sake of argument, consider the brachial and crural indices a single trait, 
since they tend to covary, and are likely influenced by the same gene complexes. 
Likewise, they are almost certainly influenced by the same environmental factors. 



GOUGH'S CAVE 1 : ASSESSMENT OF BODY SIZE AND SHAPE 



43 



the modern world, high brachial and crural indices tend to be 
associated with longer limbs. Not only have Trinkaus (1981) and 
Meadows & Jantz (1995) documented this, but Roberts' (1978) 
relative forearm index (the anthropometric equivalent of the brachial 
index) is also positively associated with temperature, and thus tends 
to be found in absolutely longer-limbed groups. However, while the 
association between these indices and limb length is a very real one, 
there remains much variability in these features (Holliday, 1999). 
For example, among the global sample of recent humans used for 
this analysis, correlations between the brachial index and total arm 
length (humeral length + radius length), and between the crural 
index and total lower limb length (femoral + tibial length) are 
significant, but are not particularly high (for the former relationship, 
r = 0.12, p = 0.0036, n = 63 1 ; for the latter, r = 0. 15, p = 0.0001 , n = 
680). Thus, while there is a clear tendency among recent humans for 
brachial and crural indices to increase with overall limb length, there 
is also considerable variability in limb length, and how that length is 
distributed between the proximal and distal segments (and see 
Holliday & Ruff, 2001). As a result, there is much overlap in distal 
limb segment length proportions among individuals from broad 
geographic regions (Holliday, 1999). 

Nevertheless, when the brachial indices of recent Europeans are 
compared to Mesolithic and Late Upper Paleolithic samples, two- 
tailed t tests detect significant differences between the recent and 
fossil Europeans (Mesolithic vs. Recent, p = 0.004; LUP vs. Recent, 
p = 0.0002). The crural index yields similarly significant differences 
(Mesolithic vs. Recent, p = 0.01; LUP vs. Recent, p < 0.0001). It is 
difficult to imagine that these differences are due to mere sampling 
error in the fossil record. 

Thus, we are faced with a dichotomy. In multivariate analyses of 
shape, Mesolithic and LUP samples (unlike their EUP forebears) 
cluster among recent Europeans, yet their brachial and crural indices 
are significantly higher. Importantly, however, this does not mean 
that their limbs are long. In fact, while brachial and crural indices 
remained elevated from the EUP through the Mesolithic, total limb 
length reduce d (Frayer, 1980, 1981, 1984, 1992; Jacobs, 1983, 1985; 
Holliday, 1995, 1999). 

What, therefore, do the high brachial and crural indices of the Late 
Upper Paleolithic and Mesolithic humans, including Gough's Cave 
1, mean? As argued in Holliday (1999), this is a clear example of 
mosaic evolution. It seems likely that climatic selection due to the 
glacial cold of Europe modified what had been a more tropically- 
adapted physique into a more cold-adapted one. Yet selection acted 
more or less equally on both the proximal and distal limb segments, 
leaving the later humans with shorter limbs (and thus better adapted 
to the cold), but permitting them to retain their relatively long distal 
limb segments 2 . 

Whether some other type of selection was maintaining these high 
ratios in spite of overall reduction in limb length, or whether they 
were selectively neutral is uncertain. The most likely conclusion is 
that the brachial and crural indices are genetic markers linking the 
Late Upper Paleolithic and Mesolithic populations to their 'Cro- 
Magnon' forebears. The logical extension of this argument is that 
contra Frayer (1992) and Frayer et al. (1993), these indices are, as 
Trinkaus ( 1981) first posited, indicative of African genes in the early 
modern Europeans. 

In sum, while the total morphological pattern of the Cheddar 
Man's body proportions is European-like, it is those features for 
which he differs from the modern European condition that are of the 



2 Al least over the time period observed - at some point, apparently subsequent to the 
Mesolithic, European brachial and crural indices decreased to approximate the con- 
dition seen today. 



most interest. Specifically, it seems likely that his high brachial, 
crural indices, and TL/STH indices, reflective of relatively longer 
distal limb segments, are a retention from an earlier, largely African 
gene pool - a retention no longer seen in Europe today. 



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Bull. nat. Hist. Mus. Lond. (Geol.) 58(supp): 45-50 



Issued 26 June 2003 



Gough's Cave 1 (Somerset, England): an 
Assessment of the Sex and Age at Death 

ERIKTRINKAUS 

Department of Anthropology. Campus Box 1114, Washington University, St. Louis, MO 63130, USA 

LOUISE HUMPHREY & CHRIS STRINGER 

Department of Palaeontology, Natural History Museum, Cromwell Road, London SW7 5BD, U.K. 

STEVEN E. CHURCHILL 

Department of Biological Anthropology and Anatomy, Box 3170, Duke University Medical Center, Durham NC 
27710, USA 

ROBERT G. TAGUE 

Department of Geography and Anthropology, Louisiana State University, Baton Rouge, LA 70803, USA 

SYNOPSIS. The overall impression of the sexually dimorphic characteristics of Gough's Cave 1 is that the remains are those of 
a male. However, the specimen does present some 'female' features in the facial skeleton, the ischiopubic rami and pelvic 
apertures, combined with relatively small overall size, and an ambiguous greater sciatic notch morphology. Nevertheless, the 
various features employed for sexual diagnosis of Gough's Cave are predominantly those which indicate or strongly suggest that 
it is male, but this must be accompanied with the caveat that either this individual falls at the feminine end of the male range of 
variation or that the patterns of skeletal sexual dimorphism of the population from which it derived were modestly different from 
those of the mostly European and European-derived reference samples used for this assessment. In contrast to the ambiguities of 
sex determination for Gough's Cave 1 , the various indicators of his age-at-death are highly consistent. All of them agree in placing 
Gough's Cave 1 between his late second decade and middle third decade. He was unlikely to have been younger than about 18 
years, and most likely was not older than about 23 years at death. 



INTRODUCTION 



The remains of Gough's Cave 1 have been considered to be those of 
a 'young adult male' since Seligman & Parsons' (1914) original 
partial description of the remains (e.g., Oakley, 1971; Stringer, 
1985). The original assessment of the age in the second half of the 
third decade was based on their observations of cranial sutures, 
postcranial epiphyses and dental attrition. The sex assessment was 
based entirely on comparisons of femoral proximal and distal epi- 
physeal dimensions to those of Medieval British remains. However, 
since the remains contain many more indicators of both sex and age, 
these need to be reconsidered. 



SEX DETERMINATION 



Overall Body Size 

Overall body size can provide a good indication of sex, if the 
individual in question falls above or below the area of overlap 
between the sexes. For this, the two best indicators are femoral 
length and femoral head diameter, since the former correlates closely 
with stature and the latter with body mass. 

The femoral lengths of Gough's Cave 1 (439.0 and 433.0 mm -the 
difference due largely to differences in neck-shaft angle), fall slightly 
above an overall European Mesolithic male mean (430.8 ± 19.5 mm, 
N = 35) and on either side of the mean of a European Mesolithic male 
sample without the large Muge sample (435.9 ± 20.3 mm, N = 21) 
(for comparative samples, see Trinkaus, 2003). However, the aver- 



age femoral length of Gough's Cave 1 is only 1.23 standard devia- 
tions from an overall Mesolithic female mean (407.8 ± 22.9 mm N = 
21) and only 0.90 standard deviations from the mean of a female 
sample without the Muge remains (416.2 ±21.9 mm, N = 10). 

Similarly, the sagittal femoral head diameters of Gough's Cave 1 
(47.7 & 46.3 mm) are close to Mesolithic male means (46.3 ± 2.2 
mm, N = 32; 47.0 ± 2.6 mm, N = 17 without Muge). Yet, the z-scores 
of the mean diameter (47.0 mm) relative to the female samples are 
1 .59 for the full sample (42.7 ± 2.7 mm, N = 1 5) and only 1 .06 for the 
female sample without Muge (43.7 ± 3.1, N = 8). 

Consequently, these size considerations support a male sex deter- 
mination for Gough's Cave 1, but they are not conclusive by 
themselves relative to other skeletally sexed European Mesolithic 
remains. 

The Pelvis 

The pelvic remains of Gough's Cave 1 present a mixture of male and 
female features, plus ones that are ambiguous. Yet, the overall 
impression is that of a male pelvis with some female proportions. 

The greater sciatic notches (Trinkaus, 2003: fig. 2) appear to be 
intermediate between the classic male semi-circular form and the 
more open female pattern. In addition, the right ilium, but not the left 
one, has a clear pre-auricular sulcus. 

The ischiopubic rami (Trinkaus, 2003: fig. 3) are relatively thin 
and flare ventrally along their medial margins, a generally female 
feature (see Poulhes, 1947; Phenice, 1969). Yet, the small medio- 
lateral breadth of the symphyseal body (obturator foramen margin to 
symphyseal surface), the thickness of the superior pubic ramus, the 
absence of a subpubic concavity, and the vertically elongated shape 



© The Natural History Museum, 2003 



46 



E. TRINKAUS ETAL. 



Table 1 Results of pelvic discriminant function analysis for Gough's Cave 1 . 



Reference Sample 



Female N 



MaleN 



% Correctly Classified 



Gough's Cave 1 Sex Assignment 



Euroamericans 






pelvic variables 


40 


35 


pelvic and femoral variables 


39 


34 


Afroamericans 






pelvic variables 


42 


40 


pelvic and femoral variables 


41 


39 


Pooled Sample 






pelvic variables 


113 


123 


pelvic and femoral variables 


107 


116 



98.7% 
100% 

97.6% 
100% 

95.8% 
98.7% 



male 
male 

male 
female 

male 
male 



of the obturator foramen all indicate a male. This is supported by its 
subpubic angle (64°), which is very close to a recent Euroamerican 
male mean (63.7° ± 7.8°, N = 50) and well below that of a 
Euroamerican female sample (88.4° ± 8.5°, N = 50) (Tague, 1989) 
(other recent human samples exhibit similar mean angles and distri- 
butions for males and females (Tague, 1989)). 

At the same time, the shape of the pelvic inlet is exceptionally 
round (Trinkaus, 2003: fig. 4), since its dorso-ventral and transverse 
diameters are equal, providing an index of ca.100. In contrast, a 
sample of Euroamerican male pelves has a mean index of 79.0 (± 7.9, 
N = 47) and a female sample has a mean of 83.1 (± 10.0, N = 47) 
(Tague, 1989). Its outlet index of 104.2 falls between the means of 
those male and female samples ( 1 1 1 . 1 ± 14. 1 , N = 44 and 99.8 ± 1 1 .0, 
N = 46, respectively). 

Given the mixed indications of these individual sex characteristics 
of the Gough's Cave 1 pelvis, we performed a discriminant function 
analysis of a series of measurements of the pelvis and femur in order 
to resolve the sex assessment of Gough's Cave 1 . The measurements 
were selected for overall proportional coverage, preservation and 
body size indication. Those employed are: sacral ventral height and 
arc, sacral antero-cranial breadth, pelvic antero-posterior inlet, mid- 
plane and outlet diameters, bi-iliac breadth, pelvic inlet transverse 
breadth, minimum bi-acetabular breadth, bi-tuberous (outlet) breadth, 
sub-pubic angle, and maximum length and head diameter of the 
femur. The analyses were performed with just the pelvic dimensions 
and combining the pelvic and femoral dimensions. 

These measurements were compared to three samples. The first 
was of Euroamericans with documented sex, given the geographical 
origins of Gough's Cave 1. The second is of Afroamericans of 
documented sex, given the slightly linear body build of Gough's 
Cave 1 (Holliday & Churchill, 2003). The last includes the first two 
samples, plus four samples of Amerindians with skeletally deter- 
mined sex (see Tague (1989) for sample composition). The analyses 
were done first using only the modern human reference sample, and 
Gough's Cave 1 was then included to determine its affinities. 

As can be seen in Table 1, Gough's Cave 1 is assigned to the 
male sample in all but one case, when the reference sample con- 
sists of Afroamericans and the femoral variables are included in 
the analysis. This single exception is almost certainly a result of 
the slightly linear proportions of the Gough's Cave specimen com- 
bined with the relatively tall stature of the individuals in that 
reference sample. 

The Skull 

The overall impression from the Gough's Cave 1 cranium is that of 
a male. Although we do not have other crania from the same 
population for comparison, the prominence and volume of the 
mastoid processes suggest that the cranium is that of a male. A 



further indication that the cranium is male is the presence of a 
pronounced crest on each suprameatal triangle, which extends the 
zygomatic process almost as far as the parietal notch. The cranium 
has marked temporal lines with both the upper and lower temporal 
lines extending to the lambdoid suture. 

The appearance of the occipital is also that of a robust individual. 
It has marked superior and inferior nuchal lines and a well-defined 
external occipital crest. The area of the external occipital protuber- 
ance is partially obscured by sediment, but it is clearly not a prominent 
feature. 

The sexually diagnostic features of the upper facial region and 
frontal bone are ambiguous. The right side of the glabella and right 
supraorbital margin are partially obscured by a pathological lesion. 
Based on what can be seen, the glabella is only moderately prominent 
and the supraorbital ridges are not particularly well-defined. The 
supraorbital margin is of moderate thickness and sharpness. 

Relative to the overall impression from the cranium the mandi- 
ble appears to be that of a more gracile individual (for a detailed 
description of mandibular morphology, see Humphrey & Stringer, 
2002). The mental protuberance and mental tubercles are not 
particularly prominent. The gonial region is everted and the areas 
of attachment of the masseter and medial pterygoid are well 
defined. 

Buikstra and Ubelaker (1994) emphasised five aspects of cranial 
morphology that can be useful for sex determination. Each feature is 
scored on a five-point scale, with higher values representing more 
robust masculine features. A score of 1 indicating a probable female, 
a score of 5 indicating a probable male and a score of 3 indicating that 
the feature is ambiguous. The scores for the Gough's cave cranium 
for each of the five sexually diagnostic structures are: 

Robusticity of nuchal crest: 5; size of the mastoid process: 5; 
sharpness of the supraorbital margin: 2; prominence of the glabella: 
3; and projection of the mental eminence: 2-3. 

The skull therefore presents a mixture of robust and gracile 
characteristics. The most masculine features relate to the attachment 
of the nuchal and temporal musculature, while the supraorbital 
region and mandible present features that are not obviously mascu- 
line or feminine. 

The Gough's Cave 1 cranium was compared metrically to a 
sample of European Mesolithic and Late Upper Palaeolithic crania 
(Humphrey and Stringer, 2002). The crania were measured accord- 
ing to the system devised by Howells (1973). A total of 39 cranial 
measurements were made of Gough's Cave and the comparative 
sample included only crania on which the same set of measurements 
could be taken. Principal components analysis of 39 cranial dimen- 
sions suggests that Gough's Cave 1 is male (Humphrey & Stringer, 
2002). A stepwise discriminant analysis using the same comparative 
sample classifies Gough's Cave 1 as male (Humphrey & Stringer, 
2002). 



GOUGH'S CAVE 1 : ASSESSMENT OF SEX AND AGE AT DEATH 



47 



Summary Sex Assessment 

The overall impression of the sexually dimorphic characteristics of 
Gough's Cave 1 is that the remains are those of a male. This is 
supported by posterior and posterolateral cranial features, long bone 
lengths, many discrete pelvic traits and particularly discriminant 
functional analysis of the pel vis. However, the specimen also presents 
a series of features that are generally considered to be female 
characteristics, including several features of the facial skeleton, the 
ischiopubic rami and the pelvic apertures. This is combined with its 
overall size being well within Mesolithic female ranges of variation, 
and its ambiguous greater sciatic notch morphology. 

We feel that the various features employed for sexual diagnosis of 
Gough's Cave are predominantly those which indicate or strongly 
suggest that it is male, but this must be accompanied with the caveat 
that either this individual falls at the feminine end of the male range 
of variation or that the patterns of skeletal sexual dimorphism of the 
population from which it derived were modestly different from those 
of the mostly European and European-derived reference samples 
used for this assessment. 



AGE ASSESSMENT 

In their presentation of the Gough's Cave 1 remains, Seligman & 
Parsons (1914) make several references to its age at death, including 
'the sutures are open both extra- and intra-cranially, a condition 
which would make us fairly sure that that the individual was under 30 
years of age" (p. 255), 'the teeth in the lower jaw are very perfect and, 
although their possessor was probably between 24 and 28 years of 
age, show very little sign of grinding down' (p. 258), and 'a part of the 
left os innominatum has been preserved and shows that the epiphy- 
seal line for the crest of the ilium is not completely closed' (p. 261). 
'As all other available epiphyseal lines have disappeared in this 
skeleton we should say that death took place between the ages of 24 
and 28, and this is quite in harmony with the evidence of the skull' (p. 
261). Given the presence of a variety of other age indicators on the 
remains (of varying precision), a reassessment of these statements is 
in order. 

The Skull 

Parts of the basicranial region are missing so it is not possible to 
examine the area of the basi-occipital synchondrosis. However, most 
of the cranial vault sutures remain, permitting their assessment 
endocranially and ectocranially. 

The reliability of cranial suture closure for age estimation is 
debated. Nevertheless, several different systems have been devel- 
oped for estimating age at death from suture closure on the 
endocranial and ectocranial surfaces of the skull (e.g. Meindl & 
Lovejoy 1985, Perizonius 1984, Buikstra & Ubelaker 1994). Key 
et al. (1994) conducted a detailed investigation of cranial suture 
closure in 183 individuals of known age at death from the Christ 
Church, Spitalfields sample. Their study recorded the degree of 
closure at 54 different sites on the cranial vault. Key et al. (1994) 
demonstrated a high level of variability in suture closure with age 
in the Spitalfields sample. In particular, their study warned that 
open ectocranial sutures were found to occur with equal frequency 
at all ages, and should not be used as an indication of young age. 

The degree of suture closure in Gough's Cave 1 was evaluated 
using the methods described by Key et al. (1994). Observations 
could be made at 24/36 ectocranial sites and at 14/18 endocranial 



Table 2 Degrees of occlusal attrition in Gough's Cave 1, scored 
following the system of Molnar (1971). 

Right 



Left 



Maxilla 



Mandible 



M 1 
M 2 
M 3 

I, 
h 

c, 

P 3 
P 4 

M, 

M 2 

M, 



sites. All except two of the recording positions could be scored on 
either the left or right side of the skull. Suture closure was scored as 
at each of the sites examined. The conclusions of Key etal.{\ 994) 
suggest that this result does not provide any definitive evidence of 
age at death, and perhaps all that can be concluded in relation to the 
evidence from cranial suture closure in Gough's Cave 1 is that it does 
not conflict with other morphological indicators of a young age at 
death. 



The Dentition 

All of the teeth present in the upper and lower jaws are fully emerged 
into the tooth row, suggesting a minimum age at death of about 17 
years (Smith 1 99 1 , table 1 ). Radiographs reveal that the roots of the 
mandibular third molars are complete and appear to be completely 
closed at the apex. The mean age of attainment of apical closure of 
the third molar in a recent North American sample is 20 years for 
males and 20.7 years for females (data from Moorrees et al. 1963, 
presented by Smith 1991). The minimum age of attainment of this 
stage is just over 16 years (mean - 2sd, for age of closure of distal 
root apex (Moorrees et al, 1963). 

It is also possible to assess the degree of wear as a general 
indication of age-at-death. The occlusal attrition scores, following 
Molnar (1971), are in Table 2. In this, 1 indicates an essentially 
unworn tooth, and 3 (the highest score for Gough's Cave 1) indi- 
cates that the cusp pattern is partially or completely obliterated 
and there are small dentine patches exposed. As can be seen, all of 
the anterior teeth and three of the first molars exhibit wear stage 3, 
the third molars exhibit wear stage 1 , and the remaining teeth are 
in between. 

Of particular relevance is the amount of wear on the third molars, 
which is minimal. Both mandibular third molars have slightly pol- 
ished enamel, and there is a small wear facet on the mesio-buccal 
cusp of the right one. The difference in the amount of wear between 
the left and right teeth is consistent with the amount of wear on the 
other molars, which is higher on the right side than on the left. There 
is slight polishing on the upper third molars and a small wear facet on 
the mesio-lingual cusp of the upper right third molar. The amount of 
wear suggests that death occurred not long after the third molars 
came into occlusion. The evidence from the third molars is consist- 
ent with the relatively low level of wear on the first and second 
molars. Application of the Miles method (Miles 1978) for ageing 
using attrition on the mandibular molars indicates an age at death of 
between 1 8 and 24 years, with an age at the lower end of the scale 
being more likely. 



48 



E. TRINKAUS ETAL. 



The Axial Skeleton 

The Vertebral Column 

The indications of age-at-death in the vertebral column, as preserved 

and as observable given the partial articulation of the remains 

(originally for museum display), are as follows: 

C6 or 7: Posterior tubercle of spinous process unfused. 

Tl: Posterior tubercle unfused. 

T2 or 3: Posterior tubercle unfused. 

Til: Posterior tubercle appears to be unfused. 

T12: Posterior tubercle unfused, annular ring of the inferior surface 
is not fully fused. 

LI: Posterior tubercle appears to be unfused. 

L2: The tubercle of the spinous process is fused but the epiphyseal 
line is still open along its superior margin. The epiphyseal line 
between the secondary center of ossification of the inferior annu- 
lar ring and the centrum is also evident (but is mostly closed and 
was undergoing obliteration at the time of death). 

L3 : The tip of the spinous process is fused but the epiphyseal line is 
still open along its superior edge. The inferior and superior 
annular rings appear to be fully fused to the centrum, with the 
epiphyseal lines completely obliterated. 

S 1-S2: Between the S 1 and S2, the ventral bodies are fully separate, 
with a maximum gap between them of 2.3mm. Laterally and 
dorsally they remain unfused but the bone surfaces are in contact 
with each other. 

S2-S3: There is clear contact but no evidence of fusion between S2 
and S3 bodies. 

S3-S4: There is clear contact but no evidence of fusion between S3 
and S4 bodies. 

S4-S5: The S4 and S5 bodies are fully fused, but the line between 
them is readily apparent. 

S5-Cxl : There is no evidence of any bridging between the S5 and 
Cxi bodies. 

Summary. The secondary center of ossification for the inferior 
annular ring of the twelfth thoracic vertebra is unfused, and the 
inferior annular ring of the second lumbar vertebra is fused but the 
epiphyseal line remains visible. The superior and inferior annular 
rings of the third lumbar vertebra are clearly fused, and the epiphy- 
seal lines are obliterated. Post-mortem damage to the bones and the 
presence of reconstructive materials and adherent matrix make the 
evaluation of the developmental state of the other vertebrae difficult. 
The dorsal tubercles of the spinous processes of the sixth cervical, 
first, second, eleventh and twelfth thoracic and first lumbar vertebrae 
are clearly unfused, suggesting a relatively young age at death for 
this individual. 

Secondary centers of ossification in the vertebrae appear around 
puberty, and with the exception of the epiphyseal rings of the 
centra, are usually fused by the age of 18 years (Steele & 
Bramblett, 1988). Maturation of the annular rings usually begins 
prior to age 17 and is complete by the age of 25 (Steele & 
Bramblett, 1988). However, a considerable amount of individual 
variation exists in ages of fusion of the annular rings and other 
secondary centers in the vertebrae (McKern & Stewart, 1957). 
None of the preserved vertebrae shows any signs of osteophyte 
development, arthritis to the articular surfaces, or Schmorl's nodes 
(on the centra that can be examined), consistent with the death of 
this individual during the third decade. 

The pattern and degree of fusion of the sacral vertebral bodies is 
normal for a young adult, and by reference to Euroamerican males 
indicates an age-at-death in the mid twenties (McKern & Stewart, 
1957). 



The Costal Skeleton 

4 Right: The surface of the head is rough and irregular, likely 
representing the subchondral surface of the unfused secondary 
center of ossification for the head. 

5 Right and Left: The secondary centers of ossification for the heads 

are only partially fused (and portions are missing). 

6 Right and Left: The heads are incompletely fused and portions of 

them are missing. 

7 Left: The secondary center of ossification for the head of the left 

rib is unfused and missing. 

8 Right: The head is unfused and missing. 

9 Right and Left: The centers of ossification for the heads are 
unfused and missing. 

1 1 Right and Left: The heads are unfused and missing. 

12 Right: The head appears to be unfused. 

Summary. Most of the ribs preserving the proximal end have 
unfused or partially fused heads. The secondary centers of ossifica- 
tion for the articular tubercles are, without exception, fully fused in 
all the ribs retaining this region. Secondary centers for the head and 
tubercle generally appear around puberty and fuse between the ages 
of 1 8 and 24 (McKern & Stewart, 1 957 ), beginning in the upper and 
lower end ribs and progressing towards the middle. Apparently the 
articular tubercles followed a more accelerated schedule of fusion 
than the rib heads in the Gough's Cave 1 skeleton. The developmen- 
tal state of Cheddar Man's ribs suggests that he died in his late teens 
or early in his third decade. 

The Upper Limbs 

No degenerative changes are evident in any of the preserved upper 
limb articular surfaces, and all of the age-at-death indications are 
associated with the fusion and obliteration of the epiphyseal lines. 

The Claviculae 

Both lack the sternal epiphysis but have well preserved metaphyseal 
surfaces, making it clear that the sternal secondary centers of ossifi- 
cation were unfused. 

The Right Scapula 

All of the observable secondary centers of ossification are fully 
fused, and the epiphyseal lines are obliterated. These include the 
subcoracoid center, the infraglenoid center, the acromial center, and 
the vertebral border center (at least at the root of the spine - the only 
place this center can be evaluated). It is possible that the vertebral 
border - inferior angle center of ossification was not fully fused 
along its entire length, and that the preserved portion of the inferior 
angle represents an epiphyseal surface. Reconstructive materials 
obscure observation of the inferior angle, making evaluation of the 
state of fusion of the growth center difficult. 

The Humeri 

There is a very slight trace of an epiphyseal line on the anterior and 
medial surfaces of the proximal metaphysis just below the lesser 
tubercle and the articular surface of the head. The line is more 
apparent on the right humerus than on the left. On both humeri the 
line is largely obliterated on the dorsal and lateral surfaces. Even 
though the line is visible, the head is fully fused and the lines are near 
obliteration. Radiographically, a faint sclerotic line can be made out 
between the metaphysis and proximal epiphysis on the right hu- 
merus (despite considerable trabecular radio-opacity in the area). No 
such line can be distinguished amongst the trabeculae on the left 



GOUGH'S CAVE 1 : ASSESSMENT OF SEX AND AGE AT DEATH 



49 



side. Distally, the epiphyses and the medial epicondyle secondary 
centers of ossification are fully fused and the lines obliterated (both 
on gross and radiographic examination) on both humeri. 

The Ulnae 

The proximal and distal epiphyses are fully fused. The lines between 
the olecranon secondary centers and the proximal shafts are obliter- 
ated (radiographically as well as on gross external examination). The 
left ulna has a closed but still (barely ) visible epiphyseal line between 
the head and shaft. Sclerotic epiphyseal lines can be seen between 
the metaphyses and distal epiphyses of both ulnae, albeit more 
distinctly in the left ulna. The distal epiphyseal lines are more 
distinct in the antero-posterior than in the medio-lateral radiographs. 

The Radii 

The proximal (right and left) and distal (left) epiphyses are fully 
fused, and the epiphyseal lines are obliterated on gross examination. 
Radiographically, radiotranslucent lines can be faintly discerned on 
the right proximal and left distal radius in antero-posterior view. 

The Hand Remains 

The metacarpals and phalanges all exhibit complete fusion of the 
epiphyses externally. 

Summary 

With the exception of the proximal clavicles, all of the upper limb 
epiphyseal lines are either entirely fused and completely obliterated, 
or are essentially fused but still show a slight trace (mainly 
radiographically) of the fusion line. These age indicators are all in 
agreement, given normal variation, in assigning an age-at-death 
between approximately 1 8 and 25 years, with the absence of fusion 
in the proximal clavicle suggesting a maximum age estimate closer 
to 22 or 23 years (McKern & Stewart, 1957). 

The Lower Limbs 

The Pelvis 

The symphyseal and auricular surfaces of the Gough's Cave 1 pelvis 
are completely obscured by its articulated state, so that the age- 
related metamorphosis of these surfaces cannot be employed for age 
assessment. However, the ilium and the ischium show clear age 
indicators. On the left ischium, the tuberosity epiphysis is unfused 
along the external margin from the middle of the tuberosity to the 
medial (pubic) end of the tuberosity; internally and proximally it is 
fully fused. On the right tuberosity, there is only a hint of a persistent 
fusion line externally, but it is partially obscured by matrix. Along 
the iliac crests, there is also partial fusion of the epiphyseal lines. On 
the right side, the crest is incompletely fused from the iliac pillar to 
the iliac tuberosity, being completely unfused near the pillar and 
tuberosity and partially fused between them. On the left side, the 
crest is unfused (and absent) from the iliac pillar to the ventral 
margin of the iliac tuberosity, and then partially fused along the 
tuberosity. Ventrally, there is a fusion line still apparent externally 
(but not internally) from the anterior superior iliac spine to the region 
of the pillar. 

The Femora 

There is no trace of the epiphyseal fusion lines on the femora 
externally, for the head, trochanters or condyles. Radiographically, 
the fusion lines are completely obliterated through the trabeculae for 



the heads, the greater trochanters and the right distal epiphysis. 
However, there is a hint of a line from the middle of the condyles to 
the epicondyles in the antero-posterior radiograph of the left distal 
femur. 

The Tibiae and Fibulae 

The right tibia and fibula also show no trace externally of fusion lines 
distinct from normal capsular attachment areas around their epiphy- 
ses. Distally, both have no trace of a fusion line radiographically, but 
proximally both of the these bones show a slight indication of the 
former fusion line. In the tibia, there is a hint of a condylar fusion line 
in antero-posterior view, and the trabeculae of the proximal fibula 
exhibit radiographically a head fusion line that is largely obliterated. 

The Pedal Remains 

The two lateral metatarsals have no retention of their head epiphy- 
seal fusion lines, but the proximal metatarsal 1 still retains a slight 
indication of the base fusion line. It is apparent only in the medio- 
lateral radiographic view along the dorso-plantar middle third of the 
base. 

Summary 

The leg bone and pedal epiphyseal fusion, all of which is normally 
complete by late in the second decade, primarily indicates that this 
individual was no younger than the late second decade but is unlikely 
to be much older given the persistence of fusion lines radiographically 
around the knee and in the proximal metatarsal 1. The degree of 
fusion of the iliac crest, stages 1/2 of McKern & Stewart (1957), 
places Gough's Cave 1 most likely between the ages of 17 and 19 
with the possibility of being as old as 22. The partial fusion of the 
ischial tuberosity suggests a similar age, most likely between 17 and 
21 but unlikely to be older than about 22 years. 

Summary Age Assessment 

In contrast to the ambiguities of sexual determination of Gough's 
Cave 1, the various indicators of his age-at-death are highly consist- 
ent. All of them agree in placing Gough's Cave 1 between his late 
second decade and middle third decade. In this, the dentition sug- 
gests an age between about 1 8 and 24 years, the vertebrae suggest an 
age in the middle of the third decade, whereas the rib and appendicu- 
lar epiphyses (including the pelvis) suggest an age between the late 
second and the early third decade. It therefore appears that Gough's 
Cave 1 was unlikely to have been younger than about 1 8 years at 
death, and most likely was not older than about 23 years at death. 



REFERENCES 

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skeletal remains. Arkansas Archaeological Survey Report, 44. 
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assessment of body size and shape. Bulletin of the Natural History Museum, Geology, 

58( supplement): 37-44. 
Howells, W.W. 1973. Cranial Variation in Man. Papers of the Peabody Museum of 

Archaeology and Ethnology, 67: 1-259. 
Humphrey, L. T. & Stringer, C. 2002. The human cranial remains from Gough's Cave 

(Somerset. England). Bulletin of the Natural History Museum, Geology, 58: 153— 

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Key, C.A., Aiello, L.C. & Molleson, T. 1994. Cranial suture closure and its implica- 
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McKern, T.W. & Stewart, T.D. 1 957. Skeletal Age Changes in Young American Males. 

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E. TRINKAUS ETAL. 



Meindl, R.S. & Lovcjoy, CO. 1985. Ectocranial suture closure: a revised method for 

the determination of skeletal age based on the lateral-anterior sutures. American 

Journal of Physical Anthropology, 68: 47-56. 
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ten permanent teeth. Journal of Dental Research, 42: 1490-1502. 
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Perizonius, W.R.K. 1984. Closing and non-closing sutures in 256 crania of known age 

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Journal of Physical Anthropology, 30: 297-301. 



Poulhes, M.J. 1947. La branche ischio-pubienne: ses caracteres sexuelles. Bulletin et 

Memoires de la Societe d Anthropologic de Paris. (9) 6: 191-201. 
Seligman, C.G. & Parsons, F.G. 1914. The Cheddar Man: A skeleton of Late 

Palaeolithic date. Journal of the Royal Anthropological Institute, 44: 241-263. 
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New York. 
Steele, D.G. & Bramblett, C.A. 1988. The Anatomy and Biology of the Human 

Skeleton. College Station, Texas. 
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University of Bristol Spelaeological Society, 17: 145-152. 
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Bull. not. Hist. Mus. Land. (Geol.) 58(supp): 51-58 



Issued 26 June 2003 



Gough's Cave, Cheddar, Somerset: 
Microstratigraphy of the Late Pleistocene/ 
earliest Holocene sediments 



RICHARD I. MACPHAIL 

Institute of Archaeology, University College London, 31-34, Gordon Sq., London, WC1H OPY, UK 

PAUL GOLDBERG 

Department of Archaeology, Boston University, 675, Commonwealth Ave., Boston, Mass. 02215, USA 

SYNOPSIS. Eleven thin sections of Late-glacial and early Holocene sediments from Gough's Cave were investigated by soil 
micromorphology in order to complement analyses of contemporary faunal and human remains. Despite the paucity of continuous 
vertical and lateral stratigraphic sequences, which were the result of cave exploitation during the first half of the twentieth century, 
we were able to elucidate site formation processes relating to both Late-Glacial environmental conditions and the burial 
environment affecting human remains. 



INTRODUCTION 



During 1987-1989 the Late Pleistocene (c. 12 ka bp) to earliest 
Holocene cave sediments at Gough's Cave, Cheddar, Somerset, were 
studied in conjunction with archaeological, human bone and faunal 
studies by R. Jacobi, A. Currant, and C. Stringer (Natural History 
Museum)(Jacobi, 1985, 1991; Currant et al., 1989; Stringer, 1990, 
2000; Currant, 1991). Sedimentological investigations, like the ex- 
cavations, suffered from having only relict and fragmentary deposits 
to study, on account of the general removal of most of the cave fill 
during the opening up of the cavern during the first part of the 
twentieth century (Donovan, 1955). We therefore focused our atten- 
tion upon extant sediment sequences dispersed within the upper part 
of the cave with Late Pleistocene deposits: i) Areas I and III of the 
North Wall of the cave, ii) the 'Skeleton Rift', iii) a cemented, early 
Holocene stalagmite on the 'South Wall' , and iv ) an earliest Holocene 
sequence in the 'Sand Hole'. 



METHODS 



Field 

Undisturbed samples were collected during the excavations from 
North Wall Areas I (samples 44 and 59, G and H) and III (sample E), 
the 'Skeleton Rift' (sample D), cemented (Holocene) stalagmite on 
the 'South Wall' (sample F), and an earliest Holocene sequence in the 
'Sand Hole' (samples A, B and C)(Table 1; Fig. 1). Samples were 
impregnated with an epoxy resin and manufactured into ~8 x 6 cm 
size thin sections at the Natural History Museum, London. 

Eleven thin sections were made from Gough's Cave and were 
described according to Bullock etal. ( 1 985) and Courty etal. (1989). 
They were viewed at a number of magnifications ranging from x 1 , up 
to x400 under a polarising microscope, employing plane polarised 
light (PPL), crossed polarised light (XPL), oblique incident light 
(OIL), and ultra-violet (blue) light (UVL) (cf. Stoops, 1996). The 
combined use of these different types of illumination permit a large 
number of identifications, such as apatite (bone, guano and coprolites) 
which autofluoresce under UVL. The authors also made use of 
comparative material of Pleistocene cave sediments (e.g., Courty et 



al., 1989), including nearby Middle Pleistocene Westbury-sub- 
Mendip (Somerset) and Late Pleistocene King Arthur's Cave in the 
Wye Valley (ApSimon et al., 1992; Macphail and Goldberg, 1999). 
In addition, the number of soil micromorphological investigations of 
palaeosols dating to the Dimlington Stadial, Windermere Interstadial 
and Loch Lomond Stadial has increased greatly since this original 
work was done at Gough's Cave. These include a number of chalky 
colluvial Aller0d palaeosols (Rendzinas) from Kent (Macphail and 
Scaife 1987, Fig. 2.4;Preecee?a/., 1995), a ranker from West Sussex 
(Macphail 1995) and a palaeosol formed in scree outside King 
Arthur's Cave (Macphail et al. 1999). 



RESULTS AND INTERPRETATIONS 

Soil micromorphological descriptions and findings are summarised 
in Table 1 and illustrated in Figs 1-7. In order to simplify presenta- 
tion of the findings we have grouped the results and associated 
interpretations. 

Pleistocene Deposits 

Pleistocene deposits overlying the widespread, unfossiliferous, ba- 
sal conglomerate are composed of gravels overlain by silt-rich 
sediments (Fig. 1 ) that represent an identifiable depositional/post- 
depositional sequence. We can broadly refine these characterisations 
as follows: 1) bedded silts, sands and gravels; 2) the formation of 
banded fabrics with associated link cappings; and 3) reworked and 
disrupted silts and sands; 4) minor biological reworking by roots and 
fauna, and 5) inwashing of silts and dusty clay. 

Bedded silts (Fig. 2), sands and gravels (Fig. 4) are broadly related 
to an upward fining sequence associated with phreatic flow within 
the main chamber of the cave (cf. Gillieson, 1996, Fig. 5.3). These 
depositional episodes are tied to fluctuating/diminishing water flow 
events within the overall karstic system that give rise to a sequence 
of cobbles (e.g., the basal conglomerate; not sampled here), gravels 
(sample 59, Table 1 ), sand (samples G, H, D, E, I and 59), and mud 
(samples B and C). Very thick phreatic sands and gravels also typify 
the Middle Pleistocene basal fill at Westbury-sub-Mendip Cave 
(Macphail and Goldberg, 1999) and comparable karstic settings 
(e.g., Goldberg and Sherwood, 1994). 



© The Natural History Museum. 2003 



52 



R.I. MACPHAIL AND P. GOLDBERG 











■^JV' 






' tJ& 
























.''."' . , . ; ' \ ••-■-- ■ ? .<i :i 








» -•. 








-:• ' ". ""' ./'. ,*-,-^<; " 4C* 
















i '•"• 1 








r- 


- 








MkJn^IH 








"HI 
















"■ 








j2g^ 






•*2 


n 




, 




^HP^ffiSnMflHfll^^BM^9 








■ 






^ 


^^^^^^HM 






■■■ --A 


■ 



Fig. 1 Field photo of Gough's Cave, Area 3. lower red silts, sample I. Note faint traces of bedding next to the sampling box (8 x 6.5 cm); cf. Fig. 2. 



At Gough's Cave, the basic sedimentary sequence has been modi- 
fied by a number of post-depositional processes to produce different 
micro-sedimentary fabrics. For example, the banded fabrics/link 
capping features (Table 1 ) are the typical result of ice lensing 
produced by alternate freezing and thawing ( Romans and Robertson, 





Fig. 2 Macrophotograph of whole thin section of sample I (cf . Fig. 1 ). 
Shown here are interbedded beds of elutriated silts (Si) and clay (C). 
Width of photo is 6.5 cm. 



1974; van Vliet-Lanoe, 1985, 1986). Extreme modification by 
freezing and thawing results in the fragmentation and chaotic mixing 
of the beds and link capping features, and infilling with impure clay 
and silt (Macphail, 1999) (Fig. 4). 

Biological activity is also recorded at Gough's Cave, forming 
channels and vughs through likely rooting and faunal burrowing. 
Finally, in this sequence many of these voids have been coated with 
dusty clay that implies renewed fluid transport vertically through the 
sediments (see below). 

Sand Hole (Pleistocene to earliest Holocene) 

Here the sequence commences with the deposition of cave muds 
(sample C) that accumulated in the base of the Sand Hole. These 
muds are composed of clay with clasts of redeposited clay (Fig. 5) 
and accumulated under conditions of low energy ponding. These 
deposits have reticulate b-fabrics induced by minor shrinking and 
swelling that reflect alternating periods of wetting and drying. It is 
possible that these muds are the finest deposits within the cave 
system, recording the end member of the upward fining sequence 
present in the main chamber. It is likely that this red clay owes its 
ultimate origin to the weathering of the Carboniferous Limestone, 
and is a form of transported Beta B clay (Duchaufour, 1977). 

In the Sand Hole, the sequence continues with the 'Laminated 
Stalagmite' and the 'Frog Earth' (Figs 6, 7). The laminated stalag- 
mite is composed of cryoclastically produced fallen limestone clasts 
and clay beds, which are both partially cemented by micrite originat- 
ing from drip. These deposits are succeeded by muds containing 
large numbers of frog bones that appear to be typical of early 
Holocene faunas (Currant, NHM, pers. comm.). 



GOUGH'S CAVE: MICROSTRATIGRAPHY OF LATE PLEISTOCENE/EARLIEST HOLOCENE SEDIMENTS 



53 




Fig. 3 Gough's Sample D (Table 1 ), consisting of rounded bone (B) and clayey sediment aggregates ( Ag) in a calcareous silty clay. Note secondary 
porosity (V) composed of channels and vughs that feature secondary calcite carbonate growth. XPL; width of photo is ca. 5.4 mm. 



DISCUSSION 



It has not been an easy task to reconstruct the sedimentary history of 
Gough's Cave, because the micro-sedimentary evidence by neces- 
sity, has been gathered from the small (max. 40 mm thick) pockets of 
sediment that remain on the extreme edges of the main cave, and the 
mainly early Holocene sequence in the Sand Hole. 

The sequence at Gough's is quite localized and built up from non- 
continuous exposures within the cave, and so must be considered as 
yielding only a partial history of the cave. The sedimentary sequence 
commences with the deposition of the conglomerate, followed by 
sands and gravels that fine upwards to the red silts, with muds being 



restricted only to the Sand Hole (Table 1 ). This Late Glacial accumu- 
lation lasted from about 12,000 to 10,500 l4 C years bp, and seems to 
be roughly correlated with the Windermere Interstadial (ca. 13,000 
to 1 1 ,000 l4 C years bp) through to the Loch Lomond Stadial ( 1 1 ,000 
to 10,000 14 C years bp). Human skeletal remains occurred within the 
red silts and were variably coated with silty clay through to sand and 
fine gravel (Currant and Stringer, NHM, pers. comm.). The bones 
(Stringer, 2000) that date to ca. 13,000 to 1 1,500 radiocarbon years 
ago would thus appear to be in situ and contemporary with the lower 
energy deposition of the upward fining sequence. 

This phreatic period appears to be contemporary with both Upper 
Palaeolithic activity and the Windermere Interstadial (Table 2). The 
formation of the conglomerate can perhaps be best related to 




Fig. 4 Sample B from Gough's Cave (Table 1 ). Macroview of laminated silts and clays over conglomerate, the basal deposit. Width of photo ca. 1.1 cm. 



54 



R.I. MACPHAIL AND P. GOLDBERG 




Fig. 5 Sample C from the sand hole, a basal clay deposit including a locally reworked clay (CC). Note the reticulate birefringent fabric which is indicative 
of minor shrinking and swelling reflecting alternate periods of wetting and drying. Width of photo ca. 5.4 mm. 



cryoclastic activity and high energy phreatic flow occurring near the 
last glacial maximum. It is likely that diminishing phreatic flow and 
the upward fining sedimentary sequences situated at the sampled 
margins of the cave, occurred from the end of the Devensian (Oldest 
Dryas) to the Windermere Interstadial (B0lling/Aller0d). This inter- 
val was contemporary with Upper Palaeolithic activity that led to the 
deposition of human skeletal remains in sediments that were once 
well-bedded (Fig. 2). It seems likely that the ice lensing activity 
noted in sample H, could be related to occasional cold conditions 
continuing into the Interstadial. The presence of humans is totally 
unrecorded at this period in the samples from Areas I and HI and the 
Skeleton Rift, but a breccia remnant ( 'reindeer stalagmite' ) from the 




just below the cave roof does include some charcoal, as seen in thin 
section. 

The mild conditions of the Windermere Interstadial are best 
recorded, albeit weakly, by biological activity producing an enhanced 
porosity pattern of channels and vughs (e.g., in samples 44 and D; 
Fig. 3). Other contemporary sites in southern England have pro- 
duced longer sedimentary sequences. For example, a number of 
chalky colluvial deposits have been described from Kent and the Isle 
of Wight (e.g. Preece et ah, 1995). These are soil-sediments, with 
biological activity and pedogenesis being recorded through slightly 
enhanced amounts of organic matter, and in places, by concentra- 
tions of earthworm granules that indicate ephemeral land surfaces 
(Preece et ai, 1995). At King Arthur's Cave, Herefordshire, a very 
thin and weakly humic soil horizon was identified through soil 
micromorphology and chemistry (Macphail et al., 1999). This soil 




Fig. 6 Sample A3 composed of laminated stalagmite (S) overlying fallen 
limestone clast (LS) with stringers of red clay (C). Width of photo is ca. 
4 cm. 



Fig. 7 Sample A (Frog Earth) in sand hole showing clay beds (C) with 
included bone (B) capped by an iron-stained stalagmite deposit (S). 
Width of photo is ca. 4 cm. 



GOUGH'S CAVE: MICROSTRATIGRAPHY OF LATE PLEISTOCENE/EARLIEST HOLOCENE SEDIMENTS 



55 



had formed in Late Devensian scree produced from limestone, and 
was itself sealed by further scree dating to the ensuing Loch Lomond 
Stadial. Thus the dominance of sedimentation over 'pedogenic' and 
post-depositional effects, as at Gough's Cave is typical of Winder- 
mere Interstadial sites. One exception is the interstadial soil at 
Westhampnett, West Sussex, where a very thin in situ humic ranker 
had formed under a likely coniferous woodland cover (Macphail, 
1995). 

At Gough's Cave the renewed cold conditions of the Windermere 
Stadial are apparently recorded in the major disruption of sedimen- 
tary bedding and previously formed banded fabrics/linked cappings. 
This cold climate produced chaotic mixing of the deposits and 
resulted in the infilling of void space with impure clay and silts and 
clays (Figs 3, 4). Some channels and vughs formed previously by 
biological activity are coated and infilled, clearly revealing that the 
period of inwashing post-dated this activity (Table 2). Some of the 
clayey sediment adhering to the skeletal remains could thus be of the 
same origin, the orientation of the bones also possibly reflecting this 
period of disruption. For example, in the well-preserved Upper 
Palaeolithic occupation cave deposits at Arene Candide, Liguria, 
Italy, once- horizontal hearths were disrupted by this (Younger 
Dryas) freezing and thawing (Macphail et al., 1994). 

At the Sand Hole, the lowermost clay deposits (Fig. 5) appear to 
have been little affected by the Loch Lomond Stadial, but rather 
seem to reflect sedimentation strongly associated with faunal activ- 
ity during the Late Pleistocene-Early Holocene transition. Typical of 
earliest Holocene deposits, speleothem formation dominated sedi- 
mentation in the Sand Hole, with the moist conditions possibly also 
favouring the contemporary amphibian fauna (Currant, NHM, pers. 
comm.). 



CONCLUSIONS 

This study of the sediment micromorphology enabled us to identify 
the sedimentary sequence that was contemporaneous with the hu- 
man occupation/Windermere Interstadial, in spite of the paucity of a 
continuous sequence of cave sediments. Specifically, we were able 
to demonstrate an upward fining sedimentary sequence, pene-con- 
temporary with both cool and mild climatic effects. The last led to 
ephemeral biological activity, and is consistent with a number of 
other contemporary, sediment-dominated sites in southern England. 
The investigation also showed that not only sediments but also the 
skeletal remains themselves were likely influenced by localised 
post-depositional processes (minor reworking, fine sediment inwash), 
dating to the cooler conditions of the Loch Lomond Stadial (?). In the 
Sand Hole, the transition between the Windermere Interstadial/Loch 
Lomond Stadial and the earliest Holocene, is recorded in the 
sediments. This detailed microstratigraphic approach exemplified 
here appears to offer the ability to extract the maximum sedimentary 
information from disparate and discontinuous deposits within a 
sedimentary system. 



REFERENCES 



ApSimon, A. M., Donovan, D. T., Scott, K. & Smart, P. L. 1 992. King Arthur's Cave, 
Whitchurch, Herefordshire: a reassessment. Proceedings of the University of Bristol 
Spelaeological Society, 19: 183-249. 

Bullock, P., Fedoroff, N., Jongerius, A., Stoops, G. & Tursina, T. 1985. Handbook for 
Soil Thin Section Description. Waine Research Publications. Wolverhampton. 

Courty, M. A., Goldberg, P. & Macphail, R.I. 1989. Soils and Micromorphology in 
Archaeology. Cambridge Manuals in Archaeology. Cambridge University Press, 
Cambridge. 

Currant, A. 1991. A Late Glacial Interstadial mammal fauna from Gough's Cave, 
Somerset, England. In. Barton, R.N., Roberts A. & Roe D. (editors), The late Glacial 
in north-west Europe: human adaptation and environmental change at the end of the 
Pleistocene. CBA Research Report 77: 48-50. 

, Jacobi, R. & Stringer, C. 1989. Excavations at Gough's Cave, Somerset 1986- 

7 . Antiquity, 63: 131-136. 

Donovan, D. 1955. The Pleistocene deposits at Gough's Cave, Cheddar, including an 
account of recent excavations. Proceedings of the University of Bristol Spelaeologi- 
cal Society,!: 76-104. 

Duchaufour, P. 1977. Pedology. George Allen & Unwin, London. 

Gillieson, D. 1996. Caves: Processes, Development, Management. Blackwell, Oxford. 

Goldberg, P. & Sherwood, S. 1994. Micromorphology of Dust Cave Sediments: some 
preliminary results. Journal of Alabama Archaeology, 40, 56-64. 

Jacobi, R. 1 985. The history and literature of Pleistocene discoveries at Gough's Cave, 
Cheddar, Somerset. Proceedings of the University of Bristol Spelaeological Society, 
17: 102-115. 

1991. The Creswellian, Creswell and Cheddar. In R.N. Barton, A. Roberts & D. 

Roe (edsj The late Glacial in north-west Europe: human adaptation and environ- 
mental change at the end of the Pleistocene. CBA Research Report, 77: 128-140. 

Macphail, R.I. 1995. Report on the soils at Westhampnett bypass, West Sussex: with 
special reference to the micromorphology of the late-glacial soil and Marl at Area 3. 
Unpublished report to Wessex Archaeology, Southampton. 

1999. Sediment micromorphology. In, Roberts M. B. & Parfitt S. A. (editors), 

Boxgrove, A Middle Pleistocene Hominid Site at Eartham Quarry, Boxgrove, West 
Sussex. English Heritage Archaeological Reports, London., 17: 1 18-148. 

, Crowther, J. & Cruise, G. M., 1999. King Arthur's Cave: soils of the Allerod 

palaeosol. Unpublished report to N. Barton, Oxford Brookes University. 

& Goldberg, P. 1 999 The Soil Micromorphological Investigation: Westbury-sub- 

Mendip, Somerset. Pp. 59-86, In, Andrews, P., Stringer, C. B. & Currant, A. (editors), 
Westbury Cave: the Natural History Museum Excavations 1976-1984. Western 
Academic & Specialist Press Ltd., Bristol. 

, Hather, J., Hillson, S. & Maggi, R. 1994. The Upper Pleistocene deposits at 

Arene Candide: Soil micromophology of some samples from the Cardini 1940-42 
Excavation. Quaternaria Nova, Rome, IV: 79-100. 

& Scaife, R.G. 1 987. The geographical and environmental background. In, Bird, 

J. & Bird, D.G (editors), The Archaeology of Surrey to 1540, pp. 31-51. Surrey 
Archaeological Society, Guildford. 

Preece, R.C., Kemp, R.A. & Hutchinson, J.N. 1995. A Late-glacial colluvial se- 
quence at Watcombe Bottom, Ventnor, Isle of Wight, England. Journal of Quaternary 
Science, 10(2): 107-121. 

Romans, J. C. C. & Robertson, L. 1974. Some aspects of the genesis of alpine and 
upland soils in the British Isles. In, Rutherford, G K. (editor), Soil Microscopy, pp. 
498-510. Limestone Press, Kingston, Ontario. 

Stoops, G. 1996. Complementary techniques for the study of thin sections of archaeo- 
logical materials. In, Castelletti, L. & Cremaschi, M. (editors), XIII International 
Congress of Prehistoric and Protohistoric Sciences Forli-Italia-8/14 September 
1996. A.B.A.C.O., Forli, pp. 175-182. 

Stringer, C. 1 990. Hominid remains - an up-date: British Isles. 40 pp. Universite Libre, 
Bruxelles 

2000. The Gough's Cave human fossils: an introduction. Bulletin of the British 

Museum (Natural History), Geology, 56: 135-139. 

Van Vliet-Lanoe, B. 1985. Frost effects in soils. Pp. 1 17-158, In, Boardman, J. (editor), 
Soils and Quaternary Landscape Evolution. John Wiley & Sons, Chichester. 

1986. Micromorphology. Pp. 91-96, In, Callow, P. & Cornford, J. M. (editors), La 

Cotte de St. Brelade, 1961-1978. Geo Books, Norwich. 



ACKNOWLEDGEMENTS. We gratefully acknowledge a grant from Natural 
History Museum Interdisciplinary Research Fund and the help of Chris Jones 
for making the thin sections; some thin sections were made at the Institut 
National Agronomique, Paris-Grignon (M. A. Courty and N. Fedoroff). The 
help and encouragement of Chris Stringer, Andy Currant and Roger Jacobi is 
very much appreciated as well as those of others of the excavation team who 
facilitated this study. We also thank the anonymous referee for their com- 
ments. 



56 



R.I. MACPHAIL AND P. GOLDBERG 



Table 1 Selected soil micromorphological observations. 



Area/Context/ 
Unit 


Sample 


Relative 
Depth (m) 


Field 


Micromorphology 


Sand Hole' 










'Frog earth' 


A upper 




0-0.19 m: 
Yellowish red 
(5YR4/6) sandy 
loam. 


Structure: Massive, intermixed coarse silts and fine sand with thin 
beds and very coarse infills of silty clay; coarse vughy porosity. Very 
abundant sand-size bone, coprolite and possible fish bones; occasional 
long (3 cm) amphibian? bones; vertebra? also present; inclusions of 
stalagmite. Microfabric: C:F, 80:20, speckled brown (PPL), medium 
to high interference colours (close porphyric, crystallitic b-fabric; 
XPL), orange brown (OIL); rare charcoal. 


'Laminated 
stalagmite' 


A lower 


0.11-0.27 


0.19-0.41 m:pink 
(5YR7/4) laminated 
stalagmite. 


Structure: Very finely (200 um) bedded stalagmite with coarse 
limestone inclusions and silty clay bands. Likely lichen/algal 
stalagmite growth. Lower part contains few very leached bone 
fragments, otherwise sterile. Microfabric: whitish to cloudy grey 
(PPL), high to very high interference colours (crystallitic b-fabric); 
white to greyish brown (OIL); very abundant pseudomorphs of plant 
material in places. Porous basal stony layer features abundant micritic 
(with clay staining) coatings. 


'Brown Clay' 


B 


0.42-0.47 


0.41-0.49 m: 
reddish brown 
(5YR5/4) clay, with 
whitish patches 
(bone ghosts?); clay 
mixed with 
stalagmite and 
blackened, possible 
reindeer bones. 


Lower part of this massive and heterogeneous clay is sterile except for 
rare plant fragments, while the upper part is rich in very fine to coarse 
sand-size, very pale bone, with 'vole' teeth. Under UVL, the bone has 
a whitish grey autofluorescence; pores within the bone are also infilled 
with dirty clay. The sediment is also characterised by a patchy 
autofluorescence under UVL, and a partially closed vughy porosity. 
Microfabric: C:F, 80:20; dominant (clast supported) angular small 
stone size limestone and calcite, common angular to subrounded fine 
to medium sand and silt-size quartz: speckled brown and greyish 
brown (PPL), moderate to high interference colours (close porphyric, 
crystallitic b-fabric; XPL), bright orange brown (OIL); frequent (15%) 
coarse packing voids, with many clayey/calcitic coatings in the lower 
part and many dark red clay coatings in upper part. 


'Red Clay' 


C; 2 thin 
sections 


0.50-0.66 


0.49-0.90+ m: 
yellowish red 
(5YR4/6) 'sterile'? 
clay over limestone 
blocks. 


Structure: Dense massive, with rare interconnecting channels, fine 
bone and occasional plant fragments and possible in situ roots. At the 
base, cracks are present alongside rare plant fragments and many 
sand-size red clay papules (Fig. 2); C:F. 20:80; silt-size quartz, and 
rare very fine sand; speckled dark yellow/reddish brown (PPL), low 
interference colours (open porphyric, reticulate b-fabric; XPL), orange 
brown (OIL). 


South Wall 










'Reindeer 
stalagmite' 


F; 2 thin 
sections 




Post stalagmite 
brown silt and 
charcoal, over 
stalagmite 
containing a 
reindeer tooth and 
charcoal. 


Structure: Massive with disrupted beds and many closed vughs. 
Poorly sorted with stone size, angular limestone and sand-size quartz; 
charcoal, many bones and occasional teeth; C:F, 60:40; common to 
dominant stone inclusions (some limestone showing etching), with 
frequent to common medium sand of quartz and flint, with very few 
fine sand size to gravel size bone - some brown stained some leached 
and rounded (ex-regurgitation pellets?), possible teeth fragments; 
occasional sand size rounded charcoal; patchy grey and brown (PPL), 
medium and high interference colours (gefuric and open porphyric, 
crystallitic b-fabric, XPL), grey and pale brown (OIL); many 1-8 mm 
thick silty clay pans (width of slide): abundant brown clayey micritic 
hypocoatings on voids and embedding large clasts. 


North Wall 










Area 1 (1987) 










Red Silt 


44 


0-0.15 


Red Silt. 


Structure: Massive to weakly coarse platy; Porosity: lower 20 mm 
-5% voids, with fine channels and vughs; uppermost 25 mm 15-20% 
voids of coarse, smooth-walled vughs and channels. Mineral: C:F, 
85:15 in lower 20 mm with horizontal fine (200 um) bands of 100:00 
(elutriated) very dominant silt to fine sand size angular quartz (and 
mica); with frequent fine, medium and coarse sand size quartz, 
limestone (also calcite); poorly rounded fragments of clay pans, 
clay/soil granules, sediment papules. Top 25 mm: dominant very 
coarse sand size quartz, quartzite, ferruginous nodules, etched calcite 



continued . 



GOUGH'S CAVE: MICROSTRATIGRAPHY OF LATE PLEISTOCENE/EARLIEST HOLOCENE SEDIMENTS 



57 



Table 1 continued 











clay /soil granules, sediment papules. Top 25 mm: dominant very 
coarse sand size quartz, quartzite, ferruginous nodules, etched calcite 
and weathered limestone. Inclusions of silt size pale yellowish 
autofluorescent material (UVL) representing reworked phosphatic 
coprolitic material. Fine mineral is composed of very dominant pale 
brown, dotted (PPL), low interference colours (close porphyric 
speckled b-fabric, XPL), pale orange brown (OIL). Very little organic 
matter. Pedofeatures: very abundant textural pedofeatures. Lower 20 
mm: very abundant intercalations, occasional very dusty clay void 
coatings: in upper 25 mm very abundant, very dusty/impure clay void 
coatings: coatings on all void surfaces and ped faces; fabric 
pedofeatures composed of abundant inclusions of banded dusty 
clay/link capping material as fragments. 


Gravels over 
conglomerate 


59 




Gravels. 


Structure: massive. Porosity: 10-20% voids generally coarse 
channels and vughs. Mineral: C:F. 85:15 (silts) to 95:5 (gravels). 
Common gravel to very coarse sand size limestone (subangular to 
angular): various rounded/weathered aragonite. with flint/chert, 
siltstone, etc. Common silt size and fine sand size quartz with very 
few mica. Fine material composed of pale brown, speckled (PPL), 
low interference colors (close porphyric. speckled b-fabric. XPL), pale 
brown orange (OIL). No obvious organic matter. Pedofeatures: very 
abundant intercalations, dusty/impure clay void coatings, pans, 
"micro-channel infills". Very abundant banded fabric of clean silts. 


Area 1 (LI 02 
metre square) 










Red Silt 


G 


0-0.095 


Red silt beneath 
cave roof. 


As Sample 44. Massive, very fine and coarse banded material with 
lamina fabric; generally closed vughs. An upward fining sequence of 
coarse silt to fine silty and clay. Contains rare plant fragments as well 
as phosphate clasts. 


Red Silt 


H 


0.33-0.40 


Red silt beneath 
cave roof, over 
conglomerate. 


As Sample 44. Massive/fine banded, composed of strongly elutriated 
coarse silts with few mixed in coarse clayey fragments and sand size 
material. Top of sample contains a gravel and silt band. 


Skeleton Rift 










Red Silt 


D 


0-0.17 


0-0.05 m: Red silt 
beneath cave roof. 
0.05+ m: 
Conglomerate. 


As Samples 44 and 59. Massive with coarse banded silts. Upwards, 
deposit is composed of gravel-rich silts with stone-sized angular to 
subangular limestone clasts. Deposit is poorly sorted, mainly silt size 
material, with common sand, strongly disrupted banded fabric of 
dusty clay and silt pans with included sand. Inclusions of likely 
fragmented link cappings as sand size clasts; rounded bone present. 


Area 3 










Red Silt 


E 


0-0.16 


0-0.16 m: Red silt 
beneath cave roof 
(concrete floor) 
over conglomerate. 


As Sample 44. Breccia; massive with few vughs and fine channels. 
Highly compact sediment; weakly to moderately impregnated with 
calcium carbonate; poorly sorted mixture of stone size, subangular 
limestone and a matrix of dominantly silt size quartz, mica and calcite. 
Also includes coarse sand size quartz, weathered limestone, fossils 
and speleothem. Patchy iron staining and depletion; possible 
occasional charcoal stuck to the roof. Few included, embedded/coated 
grains. Occasional thin to thick, very dusty to impure coatings. 


Red Silt 


I 


0-0.16 


0-0.16 m: Red silt 
beneath cave roof 
(concrete floor) 
over conglomerate. 


Massively banded silts and clays with repeated graded beds. Some 
fine channeling at the top. Most of porosity in the form of packing 
voids. 



58 



R.I. MACPHAIL AND P. GOLDBERG 



Table 2 Tentative summary of sedimentary activity in Gough's Cave as reconstructed from soil micromorhpology. 



Age ( 14 C yrs 
bp x 1000) 


Period 


Deposition 


Post-Deposition 


Comments 


9 










10 


Early Holocene 


Formation of stalagmite and 
frog bone-rich muds in the 
Sand Hole 


Stalagmite formation. 


Climatic amelioration 
associated with warm and 
moist conditions (cf. frogs). 


11-10 


Loch Lomond Stadial 
(Younger Dryas) 


Accumulation of breccia 
sediments in Sand Hole 


Patchy formation of banded 
fabric and link capping, 
accompanied in places by 
physical mixing. Washing of 
impure clay into voids. 


Renewal of cold conditions 
leading to freezing and 
thawing associated with 
increased meltwater activity. 


13 to 11 


Windermere Interstadial 
(Bolling/Allerod) 


Upward fining sequence 
from gravels and sands 
through to the Red Silt; 
deposition of mud in the 
Sand Hole 


Ephemeral biological 
activity producing channels 
and vughs. 

Localized ice lensing in the 
lowermost Red Silt. 


Increased moderation in 
climate. 

Diminishing phreatic flow, 
possibly accompanied by ice- 
lensing in the early stages 


16 to 13 


Devensian (Oldest 
Dryas) 


Formation of Conglomerate 




Cryoclastic activity 
accompanied by high energy 
phreatic flow. 



Bull. nat. Hist. Mus. bond. (Geol.) 58(supp): 59-81 



Issued 26 June 2003 



Cannibalism in Britain: Taphonomy of the 
Creswellian (Pleistocene) faunal and human 
remains from Gough's Cave (Somerset, 
England) 



P.ANDREWS 

The Natural History Museum, Cromwell Road, London SW7 5BD, UK 

Y. FERNANDEZ-JALVO 

Museo Nacional de Ciencias Naturales, 28006 Madrid, Spain 



SYNOPSIS. Human induced damage is the main taphonomic modification observed on the fossil bone assemblage of Gough's 
cave. Fossils from this site are very fragmentary, showing abundant cut-marks, percussion marks and peeling. Some specimens, 
however, are complete (ribs, vertebrae, carpal-tarsal bones and phalanges), but these elements are characterised by low marrow 
content where breakage to open the bone is not needed. Human remains recovered from this site show similar butchering patterns 
to other animals suggesting skinning, dismembering, defleshing and marrow extraction activities. Excavations during the 1986- 
1987 seasons showed that the human remains appear at the site randomly mixed with animal bones, with no specific distribution 
or arrangement of human bones. The evidence from this distribution indicates equal treatment of human and animal remains, and 
the analysis of cut-marks and other modifications suggests that both humans and animals were accumulated as the discarded food 
remains of the human population. This is interpreted as nutritional cannibalism. One exception to this is seen in the slight 
differences in skull treatment compared with other sites, suggesting a possible element of ritual cannibalism (cf Fontbregoua, the 
French Neolithic site, ca 4000 BC). 



INTRODUCTION 



Human remains from Gough's cave (Cheddar) have been recovered 
during several excavation seasons. They were found together with 
abundant remains of other vertebrate animals and stone tools from 
Oxygen Isotope Stage 2 deposits, and most come from the Late 
Pleistocene interstadial, 11,500-13,000 radiocarbon years ago 
(Stringer 2000). 

The early excavations during the late 1920's and early 1950's 
took place over a wide area of the cave, and although abundant 
fossil remains were recovered, no record was kept of the bone 
distributions. A joint excavation undertaken by the University of 
Lancaster and The Natural History Museum (UL-NHM) was much 
more restricted in extent, with most of the bones coming from 
about one cubic metre of fine gravel and silt between a large rock 
and the north wall of the cave during 1986-92 (Stringer 2000). 
These were excavated, however, with much greater precision, and 
records of the fossils and stratigraphy were kept in meticulous 
detail, so that more information is available from this small area 
than for the whole of the previous, much more extensive, excava- 
tions. In addition, fossils recovered by this recent excavation have 
been found to refit with remains recovered by the earlier, indicat- 
ing that it is the same fossil bone assemblage. The UL-NHM 
seasons have been essential in interpreting the site formation and 
the type of cannibalism practised by Homo sapiens about 12,000 
years ago. 

Cannibalism among humans has been a taboo topic and is still 
today a controversial aspect of human behaviour. By definition, a 
cannibal is a person or animal that eats any type of tissue of another 
individual of its own kind. Permissive tolerance of human cannibal- 
ism has traditionally occurred when referred to 'primitive' societies, 
but critical reviews such as Arens, (1979) have been sceptical of 



cannibalism claims based on written references or oral tradition. 
Taphonomic studies of bone remains of the victims have been the 
only way to validate some claims for cannibalism (Villa et ai, 1986a, 
1 986b; White, 1 992; Turner and Turner, 1 999; Fernandez-Jalvo et ai. 
1999;Degusta, 1999,Defleurefa/. 1999). The oldest case confirmed 
as cannibalistic practice among humans was described at the early 
Pleistocene site of Gran Dolina (TD6, Atapuerca, Burgos, Spain), 
but a recent study has discovered cut-marks on a right zygomati- 
comaxillary specimen from the Plio-Pleistocene site of Sterkfontein 
(South Africa) that may suggest an earliest case of human damage on 
human remains (Travis et al, in press). According to these authors, 
cut-marks appear on areas of ligament and muscle insertions, sug- 
gesting cuts were made on purpose to cut meat. Surprisingly this is 
the only specimen showing butchering marks, absent on the remain- 
ing 763 macro-mammalian fossil specimens, including the rest of 
the hominid remains recovered from the site. 

Cut-marks are of great significance in coming to an understanding 
of prehistoric human behaviour, but on their own they cannot be used 
as direct evidence of cannibalism. Cut-marks may appear on human 
skeletons as result of mortuary rituals, practices still current today, 
where human carcasses are defleshed but meat or marrow is not 
consumed. Cuts may be frequent on these skeletons, although canni- 
balism is absent. Sometimes carcasses are defleshed and meat or 
organs eaten as result of rituals in relation to beliefs or religion. The 
identification of nutritional cannibalism, in contrast to ritual, is 
based on a combination of indicators, the main criterion of which is 
the comparison of human and animal remains from the same ar- 
chaeological context. If a human population was living by hunting, 
and it did not distinguish between animal and human prey, the 
processing marks left on the bones of both human and animal should 
be the same. Turner (1983) has given several criteria for recognising 
nutritional cannibalism, but the most basic criteria by Villa et al. 
( 1986a, pg 43 1 ) are as follows: 



© The Natural History Museum. 2003 



60 



P. ANDREWS AND Y. FERNANDEZ-JALVO 



1. Similar butchering techniques in human and animal remains. 
Frequency, location and type of verified cut-marks and chop- 
marks on human and animal bones must be similar, allowing for 
anatomical differences between humans and animals. 

2. Similar patterns of long bone breakage that might facilitate 
marrow extraction. 

3. Identical patterns of post-processing discard of human and ani- 
mal remains. 

4. Evidence of cooking; if present, such evidence should indicate 
comparable treatment of humans and animal remains. 

Previous work on Gough's cave material has come to contradictory 
conclusions. Cook ( 1986) attributed cut-marks on human remains to 
natural damage produced by trampling, with the exception of an 
adult mandible (Gough's cave 6) that shows evidence for deliberate 
human activity related to post mortem removal of the tongue. Apart 
from this human fossil, Cook found equivocal cut-marks on animal 
bones from the site that indicates dismembering activities. Cook, 
therefore, concluded that cannibalism was absent at Gough's cave. 
In contrast to this, Currant, Jacobi and Stringer ( 1989) consider that 
there is no doubt about human processing of parts of the body at or 
close to the time of death based on the new material from the 1 987 
collections. Similarly, Charles (1998) suggests cannibalism was the 
key factor based on the intermixing of human with animal bones in 
the deposits. 

It is our intention here to show the results of a taphonomic analysis 
of Gough's Cave fossil remains. Both human and animal bones will 
be treated equally so that their modifications can be compared with 
a view to seeing if the agents responsible for the animal bones are the 
same as those responsible for the human bones. We will focus 
particularly on the evidence of cut-marks, which are present on both, 
to see if there is any difference in distribution and/or type of cut- 
marks. In addition, we will examine features of bone fracture and 
bone distribution that may contribute to the hypothesis of human 
cannibalism at the site. 



METHODS AND MATERIAL 

The fossil material here analysed consists of 240 human and other 
animal fossil bone fragments. These are in the collection of the 
Natural History Museum in London. In addition, there are a number 
of fossil bones at the local museum in Cheddar Gorge that we have 
not had the opportunity of studying and have therefore not been 
included. Both human and animal fossil bones have been examined 
with the aid of a binocular microscope. Some specimens were 
analysed using scanning electron microscopy (SEM), an ISI ABT55 
SEM-fitted with an environmental chamber, operating in the back- 
scattered electron emission mode at 20 kV, which is housed at The 
Natural History Museum (London). This type of microscope enables 
specimens to be directly analysed with no necessity for coating 
(Taylor, 1986). 

Breakage has been analysed following the method of Villa and 
Mahieu(1991): 

1. Number of fractures. 

2. Fracture angle: oblique/right/mixed (oblique and right). 

3. Fracture outline: transverse/curved- V-shaped/intermediate/ lon- 
gitudinal. 

4. Fracture edge: smooth/jagged. 

5. Shaft circumference: 1, circumference is <Vi of the original; 2, 
circumference is > ! /2of the original; 3, complete 



6. Shaft fragmentation: 1, shafts < l A of original length; 2, length 
between l A and Vi of original length; 3, length between Vi and % 
of original length; 4, length > 3 A of original length (complete). 

Unfortunately, most remains from the study collection had been 
glued together and traits of fracture angle, fracture outline and 
fracture edge could not always be identified and quantified. Other 
fracture traits such as peeling (White, 1992), percussion pits 
(Blumenschine & Selvagio. 1988), adhering flakes (White, 1992) 
and conchoidal percussion scars (Blumenschine 1 988) were recorded 
as present or absent. 

Bone surface modifications attributed to human action were iden- 
tified as tool-induced modifications such as incisions, scrape marks, 
chop-marks, hammer/anvil striations. Emplacement of cut-marks 
and identification of the muscles or tendons affected by the cuts were 
recorded. Post-depositional surface modifications were identified as 
weathering, desquamation, trampling marks, polishing, rounding, 
gnawing or tooth marks. Post-burial modifications recorded were 
manganese oxide stains, concretion (cemented sediment heavily 
attached to the fossil), soil corrosion or root-marks. 

Tooth marks were described and measured separately for all 
anatomical items following Andrews and Fernandez-Jalvo (1997): 

a. Carnivore pits on bone surface (minimum dimension) 

b. Carnivore gnawing on bone surface (transverse measurement of 
grooves) 

c. Carnivore pits on articular surfaces. 

d. Carnivore punctures on spiral breaks 

e. Carnivore punctures on transverse breaks 

f. Carnivore punctures on split shafts 

g. Multiple molar pits made by multi-cuspid teeth, 
h. Carnivore punctures on intact bone edges 



RESULTS 

The results of the taphonomic analysis are displayed in Table 1 . The 
main taphonomic modifications that affect these fossils is human 
activity as seen at this table. 

Species represented 

The Gough's cave human material consists of both crania and 
postcrania. The former indicate the presence of five individuals, two 
adults, two adolescents and one child (Stringer 2000). The adults are 
represented by a calotte, part of a second calotte and two maxillae 
and two mandibles. The adolescents are represented by a cranium, 
two maxillae and one mandible, again suggesting two individuals. 
The child has a single calvaria. Depending on how the adolescent 
material is associated, there is a minimum of five individuals in the 
Gough's Cave deposits (Stringer 2000, Humphrey & Stringer 2002). 
The taxonomic identification of the non-human collection analysed 
here has been done by A. Currant, R.M. Jacobi and C. Stringer. The 
species found are Equus ferns, Cervus elephas. Bos primigenius. Sus 
scrofa, Lepus timidus. The most abundant species represented in the 
study collection are the equids (Table 1 ), with 1 32 specimens, and this 
compares with 88 human and 42 cervids. Only two bone fragments of 
bovid, an astragalus and a tarsal, have been recovered from the study 
collection, and one fragment each of rabbit (tibia) and suid (mandible) 
species. The latter species have some impact marks, but they are too 
few to come to any conclusions about their nature and origin, and so 
our analyses here will concentrate on the modifications of the three 
common groups, one of which of course are the humans. 



CANNIBALISM IN BRITAIN: TAPHONOMY OF FAUNAL AND HUMAN REMAINS FROM GOUGH'S CAVE 



61 



Table 1 Summary of taphonomic modifications seen on the fossil bones from Gough's Cave. Modifications are shown for each major postcranial element, 
which are listed in column 1 . The total number of specimens (N) for each element is in column 2, and in column 3 the distribution of modifications by 
human action is shown for four taxonomic categories: human (h), equid (e), cervid (c) and indeterminate large mammal (m). The same distribution is 
shown for six types of modifications in the remainder of the table as explained in the text. 



Anatomical 
element 


Total 

N 


N 


Cut-marks 


Percussion 
marks 


Concoidal 
scars 


Adhered 
flakes 


Removed 
flakes 


Peeling 






h 


e 


c 


m 


h 


e 


c 


m 


h 


e 


c 


m 


h 


e 


c 


m 


h 


e 


c 


m 


h 


e 


c 


m 


h 


e 


c 


m 


Cranial 


4 


2 





2 




2 










2 
































2 










2 










Hemi-maxillae 


9 


2 


3 


4 




2 


2 


2 




2 


2 


2 


















1 





















1 







Hemi-mandible 


27 


6 


12 


9 




2 


3 


2 







9 


4 





















1 















1 


3 


1 




Hyoid 


1 





1 










1 










1 



















































Clavicle 


3 


3 










3 






















































1 










Humeri 


6 


6 










4 










3 










1 










1 










i 





















Radii 


6 


5 


1 







1 


1 







3 


1 
































1 







1 










Ulnae 


5 


4 


1 







2 


1 







1 


1 


















1 





















2 










Scapulae 


4 


4 










4 










1 











































2 










Ribs 


45 


40 






5 


20 






5 


8 






3 


































8 









Vertebrae 


19 


7 


7 


5 




3 


6 


2 




1 


1 








































2 










Pelves 


2 





2 










2 






























































Femurs 


1 


1 





















1 






















































Fibula 


1 


1 










1 
































1 
































Tibiae 


8 


1 


3 


4 







2 


3 




1 


1 


1 







1 
























1 




1 










Long bones 


3 


2 





1 




1 





1 




1 



























1 


























Patellae 


1 





1 









































































Carpo-tarsal 


35 





28 


7 







12 


3 







1 





















1 





























Metapodial 


35 


5 


24 


6 







12 


6 




1 


19 


2 







4 





















1 


1 




1 










Phalanges 


54 


6 


49 


4 







29 


2 







27 


2 
















































Totals 


269 


90 


132 


42 


5 


45 


71 


21 


5 


25 


63 


il 


3 


1 


5 








3 


2 


2 





3 


2 


2 





21 


4 


1 






Skeletal elements 

Anatomical elements of humans and other large mammals (horses 
and deer) recorded at the site suggest some differences between 
element representation. In general terms, human skeletons are better 
represented than are those of any of the other large mammals. 
Human skeletons show a relatively high abundance of cranial re- 
mains, ribs, scapulae, and arms (Table 1). In contrast, vertebrae are 
notable for their near absence, despite the abundance of ribs that 
were found in association (although not articulation) at the site. 
There is also a peculiar absence of pelves, carpo-tarsal bones and 
phalanges which are relatively abundant among horses or deer. 
Similarly, cranial elements, especially mandibles, are also abundant 
for both horses and deer, but while metapodials and phalanges are 
abundant, most limb bones are poorly represented. Horses have an 
extraordinarily high abundance of phalanges, which are not gener- 
ally common in human occupation sites. Skeletal element proportions 
are summarized in Table 2. 

Anatomical elements: limb bones 

Five upper limb bones from Gough's cave have moderately complete 
shafts, three clavicles, one humerus and two radii and ulnae. These 
were recovered in a fragmentary state but reconstructed in the 
laboratory. For example ulna M54066 is made up by six fragments 
that make up most of the right ulna (Churchill 2001) and it has a 
possible antimere in M54067. Two of the clavicles are antimeres, 
and the four scapulae have been interpreted as representing two 
males and one female individuals. All have cut-marks, sometimes 
extensive. The lower limb bones are similarly fragmentary, with no 
complete bones. There are four left femora, although only one was 



Table 2 Skeletal proportions of the anatomical elements recorded for 6 
humans, 10 equids and 6 cervids from Gough's cave. Skeletal elements 
are shown on the left, and percentage occurrences of elements based on 
numbers present (N ) divided by numbers of that element present in the 
skeleton (N ) multiplied by the MNI. 





Human 


Equid 


Cervid 






N a 


N, 




N a 


N, 




N a 


N, 


skull 


50% 


3 


1 


0% 





1 


33% 


2 


1 


hemi-maxilla 


17% 


2 


2 


15% 


3 


2 


33% 


4 


2 


hemi-mandible 


50% 


6 


2 


60% 


12 


2 


75% 


9 


2 


hyoid 


0% 





1 


10% 


1 


1 


0% 





1 


clavicle 


25% 


3 


2 


0% 








0% 








humerus 


50% 


6 


2 


0% 





2 


0% 





2 


radius 


42% 


5 


2 


5% 


1 


2 


0% 





2 


ulna 


33% 


4 


2 


5% 


1 


2 


0% 





2 


scapula 


33% 


4 


2 


0% 





2 


0% 





2 


rib 


28% 


40 


24 


0% 





36 


3% 


5 


26 


vertebra 


7% 


10 


25 


2% 


7 


32 


3% 


5 


28 


pelvis 


0% 





2 


10% 


2 


2 


0% 





2 


femur 


42% 


5 


2 


0% 





2 


0% 





2 


fibula 


8% 


1 


2 


0% 





2 


0% 





2 


tibia 


75% 


9 


2 


15% 


3 


2 


33% 


4 


2 


patella 


0% 





2 


5% 


1 


2 


0% 





2 


carpo-tarsal 


0% 





28 


11% 


28 


26 


5% 


7 


22 


metapodial 


4% 


5 


20 


60% 


24 


4 


25% 


6 


4 


phalange 


0% 


1 


56 


41% 


49 


12 


3% 


4 


24 


long bones 




2 













1 




Total number 


8% 


90 


179 


10% 


132 


134 


6% 


47 


128 


MNI 


6 


10 


6 



62 



P. ANDREWS AND Y. FERNANDEZ-JALVO 



seen, so that at least four individuals are indicated. All four left femora 
are dyaphysis fragments that represent small individuals, and in addi- 
tion there is a right proximal femur and fragments of diaphysis from a 
larger sized individual (Trinkaus 2000), so that the MNI indicated by 
the femur is five. Nine tibia fragments indicate four individuals, two 
large and two small (Trinkaus 2000), but only one was seen. 

Humerus: There are six fragments of humeri, all of them from a 
single human individual. The shafts are split longitudinally (shaft 
circumference category 1, shaft fragmentation categories 1 and 2 
according to Villa and Mahieu 1 99 1 ). The ends are absent, with only 
one split fragment of shaft near the neck of the head (GC'87, no. 12). 
Cut-marks appear on four of the six fragments of humerus (67%). 
Cuts run obliquely along the shaft clustered or isolated covering 
rugose surfaces or muscle attachments (deltoid crest, triceps inser- 
tion or brachialis muscle). One of the specimens (GC'87, no. 12) 
preserves the area near the head, and it is here where a cut runs 
transversally across the humerus on the attachment of teres minor. 
Distally in the same fragment there are also scraping marks near the 
fracture edge. The scraping marks probably resulted from the removal 
of soft tissues that could have absorbed the blow when breaking the 
bone to extract the marrow (Binford, 1981). Three of these cut- 



marked fragments of humerus also show percussion marks along the 
broken edges. Some of these fragments also have conchoidal scars, 
adhered flakes and/or removed flakes, also located on the broken 
edge. Only one fragment of humerus has weathering in stage 1 
(Behrensmeyer, 1 978) and two are affected by trampling, but none of 
them have tooth marks. 

Summary of humeri. Total 6 specimens, all human. 
Cut-marks: 4 specimens (2 on fossils from the 1987 collections) 
Percussion marks: 3 specimens (2 on fossils from the 1987 collec- 
tions) 

Conchoidal scars: 1 specimen ( 1 on fossils from the 1987 collections) 
Adhered flake: 1 specimen 
Removed flake: 1 specimen 

Ulna: There are five fragments of ulna, four of them from humans 
(2 rights, 2 lefts, 2MNI) and one from a horse. The human fragments 
of ulna consist of longitudinal splits, as seen on the humeri, but 
several fragments have been refitted so that they now form most of 
the bone circumference (3 of them have circumference category 3 
according to the classification of Villa and Mahieu, 1 99 1 ). Two of the 
ulnae have cuts on the surface, running obliquely to the length of the 




Fig. 1 A, Left human ulna GC87-209, midshaft fragment with part of the lateral aspect of the shaft. Cut-marks run obliquely across the posterior ridge (i.e. 
along the bottom of the shaft), and another concentration occurs more distally (not shown here). There is extensive peeling at the proximal end, on the left 
as shown here, and three massive percussion impact marks can be seen medially, along the upper edge of the bone as viewed here. There is also an 
adhered flake on the lateral aspect. B, Six fragments making up most of right human ulna M54066 (GC202, 243. 1 19c). Fractures are mixed, smooth, and 
fragmentation 3/4. Breakage appears to be natural with no percussion marks and no cut-marks. C, Proximal radius and ulna GC89-071&073 of Equus 
ferus. Cut-marks are seen on the olecranon process. A, x 1 .6; B, x 0.5; C, x 0.7. 



CANNIBALISM IN BRITAIN: TAPHONOMY OF FAUNAL AND HUMAN REMAINS FROM GOUGH'S CAVE 



63 



bone and probably related to the insertion of the flexor muscles. 
Percussion marks and adhered flakes have also been observed on one 
of these two damaged ulnae (GC'87 209) along the broken edge. 
Both ulnae show clear evidence of peeling, one on the proximal 
broken edge (Fig. 1 A). The other two fragments of ulna which have 
not been damaged by human action have been refitted from several 
small split shafts found several metres apart from each other, in one 
case all the fragments coming from the 1987 excavation (M54066), 
and in the other, some fragments coming from the 1987 excavation 
and refitted with old 1927 excavation fragments (M54067). The 
proximal end of the right ulna M54066 (Fig. IB) has lateral crushing 
of the head. No cut-marks or percussion marks have been distin- 
guished on either of these two ulnae. One ulnae fragment (GC'50 
420) is weathered in stage 1 or 2 and has dispersed manganese on its 
surface. 

The only horse ulna-radius (GC'87 73) has only the proximal end 
preserved (circumference category 3, length category 1, Villa and 
Mahieu 1991). The heads of both the ulna and the radius are 
extensively cut and show percussion marks (Fig. 1C) and a flake has 
been removed from the interosseous space between ulna and radius. 
Percussion marks are present on the olecranon. 

Summary of ulnae. Total 5 specimens, 4 humans, 1 equid 

Cut-marks: 3 specimens (2 human, 1 equid) (Ion fossils from the 

1987 collections) 

Percussion marks: 2 specimens (1 human, 1 equid) (1 on fossils 

from the 1987 collections) 

Adhered flakes: 1 specimen (human) (from the 1987 collections) 

Removed flakes: 1 specimen ( 1 equid) 



Peeling: 2 specimens (human) (from the 1987 collections) 

Radius: There are five fragments of all of them humans. There are 
two with proximal articulations with complete circumference, cat- 
egory 2 and 3 (Villa and Mahieu, 1991 ) and more than the half of the 
length of the bone. Specimens M54071 and GC'87 74 are refitted 
shafts (5 and 7 respectively) of split shaft fragments, open longitudi- 
nally and mostly category 1 shaft circumference. Only one of these 
radii has cut-marks (Fig. 2A). These cuts were formerly interpreted 
as decorative engraving. They appear on the lateral surface along the 
length of the bone, bordering the origin of the flexor pollicis longus, 
but there is no muscle attachment along this part of the shaft 
(between the ulna and the radius). On the SEM we could observe that 
each group of incisions is actually a single compound mark made by 
a single stroke (Fig. 2B). Directionality is the same in every set (Fig. 
2B), and it appears to be the result of filleting, removal of the muscle 
progressively along the shaft. With regard to breakage, two of the 
radii have percussion marks, which are distributed along the longitu- 
dinal broken edge. One of the radii also shows large percussion 
marks on the anterior and posterior edges. Peeling is seen on M5407 1 
on at least three joint fragments. This specimen has many percussion 
impacts mainly along broken edges, and there are at least two large 
impact scars along the anterior side and one impact pit on the 
posterior surface. 

Summary of radii. Total 6 specimens, 5 human, 1 equid 

Cut-marks: 2 specimens (1 human, 1 equid) (1 from the 1987 

collections) 

Percussion marks: 4 specimens (3 human, 1 equid) 




3&&%t&'-^ %t 




Fig. 2 A, Right human radius GC87-74. Partial diaphysis with the anterior surface preserved for most of it length. Extensive cutmarks are present on the 
lateral surface, which is the side of the shaft away from the ulna where there are no muscle attachments, but in addition there are a few cut-marks 
proximally (on the left as seen here) on the supinator insertion. These marks have been interpreted as engraving, but all of the 'groups' of incisions are 
actually compound marks made by single strokes, with consistent directionality towards the superior aspect of the shaft. This is interpreted as filleting of 
the arm muscles progressively along the shaft. B, Scanning electron micrograph of GC87-74 cut-marks. Notice that marks are made by the same stone 
tool edge and made with a sawing motion that follows the same direction for all cuts along the bone shaft. A, x 1.1. 



64 



P. ANDREWS AND Y. FERNANDEZ-JALVO 




w 










1 


^Sj 


K ^ 


r'^l 






■ 


\B 




' v v 




" 5s, v 1 


^^1F| 








b| 












Fig. 3 A, Distal articulation of tibia M50017, left distal tibia of Rangifer tarandus. This articulates with astragalus M49914, and both have matching cut- 
marks, probably for disarticulation of the foot. B, Distal tibia, no number, of Equusferus, with cut-marks on the distal turberosity and oblique break, 
circumference 3, of the shaft. C, Right distal tibia of Equusferus, GC87-4. The distal articular surface is intact. The shaft is broken with oblique fracture, 
circumference 3. There are cut-marks on the lateral ridge of the distal tubersosity and carnivore chewing just proximal to the cut-marks, but they do not 
overlap and so their relative times of occurrence are unknown. There are extensive percussion marks on all surfaces of the shaft in the region of the 
oblique break, with a conchoidal scar posteriorly (on the right side of the break as viewed here). Finally, there is a network of shallow rootmarks on the 
shaft. A, x 1 ; B, x 0.65; C, x 0.6. 



Removed flakes: 1 specimen ( 1 equid) 
Peeling: 1 specimen ( 1 human) 

Femur: We only saw one fragment of human femur, although five 
individuals are apparently represented in the collection (Trinkaus 
2000). The femur fragment we saw is a split fragment of shaft (no 
ends, circumference category 1 , length category 1 , Villa and Mahieu, 
1991) with strongly developed linea aspera. It does not have cut- 
marks but it has percussion marks. These percussion marks are along 
the linea aspera, 3 grouped together. 

Fibula: There is only one fragment, which has been identified as 



human. It is part of the shaft having a circumference category 2 and 
length category 2 (Villa and Mahieu 1 99 1 ). It has cuts near the end of 
attachment of the soleus muscle indicating dismembering activities, 
and evidence of breakage provided by an adhered flake depressed 
into the cavity. 

Tibia: There are nine human tibia fragments, but we only saw one 
fragment, plus three of equid and four of cervid. One of the cervid 
tibiae has tooth marks on the surface. They are chewing marks on 
anatomical edges (tooth marks type c following to Andrews and 
Fernandez-Jalvo, 1997) measuring 1.4 and 0.9 mm (average 1.15mm). 
The human tibia fragment is a longitudinally split section of shaft 



CANNIBALISM IN BRITAIN: TAPHONOMY OF FAUNAL AND HUMAN REMAINS FROM GOUGH'S CAVE 



65 



with no ends (circumference category 1 and length category 1). In 
contrast to this all animal tibiae have circumferences category 3 and 
length between 1 and 2 (according to Villa and Mahieu, 1991 ). It is 
apparent from this that the animal bones are preserved differently 
from the human long bones. There are no cut-marks on the human 
tibia fragment, but percussion marks are present. On the animal 
bones, cuts appear on 5 of the 7 distal ends of tibiae. The cuts are 
related to the articulation of the tibia with the tarsals, on the distal 
ends (Fig. 3A), or on the posterior and/or anterior surfaces (Fig. 3B), 
and all are related to dismembering the ankle joint. The distal end of 
another equid tibia with a small part of the shaft shows cuts on the 
lateral maleolus, probably also related to cutting the short and long 
lateral ligaments when dismembering the foot. Another distal end 
with a small part of the shaft of cervid also has cuts on the shaft, but 
this time they appear on the opposite side of the shaft from percus- 
sion marks (see below). 

With regard to fracture, the human tibia has peeling on one of the 
ends and percussion impact scars on the edge of the longitudinal 
breakage. This suggests there were several impacts on the bone to 
open the bone longitudinally and expose the marrow. One of the 
equid tibiae (GC'87-4, Fig. 3C) has a conchoidal scar on the poste- 
rior midshaft and extensive percussion marks all round the shaft. 
Two of the four cervid tibiae show percussion marks. One of them 
(M50019) has percussion marks on the opposite side of cuts, which 
suggests that the latter could be anvil marks as result of blows on the 
bone. The other (M50017) has a flake removed on the lateral side 
near the broken edge and a percussion impact mark on the plantar 
side, near the articulation (Fig. 3A). M50017 has also trampling 
marks running transversally. 

Summary of tibiae. Total 8 specimens 1 human, 3 equid, 4 cervid 

Cut-marks: 5 specimens (2 equid, 3 cervid) (1 each of cervid and 

equid from the 1987 collections) 

Percussion marks: 3 specimens (1 human, 1 equid from the 1987 

collections) 

Conchoidal scars: 1 specimen (equid from the 1987 collections) 

Removed flakes: 1 specimen (cervid) 

Peeling: 1 specimen (human from the 1987 collections) 




Fig. 4 A, Left calcaneus M50029 of Equusferus. Much of the calcis has 
been damaged by extensive percussion marks on both sides, and there is 
a cluster of cut-marks along the upper edge just posterior to he articular 
surface. B, Medial view of left astragalus of Equusferus, M49843. Cut- 
marks are present in three clusters, one on the medial edge of the medial 
condyle, the second on the medial surface of the body, both seen here, 
and the third on the upper edge of the lateral condyle. Both figures. 
x0.9. 



Long bones (indet.): There are three fragments of long bones two 
of them identified as humans and the third as cervid, though the 
anatomical elements could not be specified. One of the human split 
shafts has oblique cuts and percussion marks on the edge of the 
fracture. The cervid shaft is formed by two longitudinally split 
fragments, having several cuts running transversally along the edge, 
and an adhered flake between the joint fracture of both fragments. 

Anatomical elements: hands and feet 

Calcaneus: There are eight calcanei, seven of them are from 
equids and only one of cervid. Cut-marks are present on five speci- 
mens (4 equids and 1 cervid): 
4 on the lateral side of the calcaneus along the plantar and the dorsal 

surfaces (4 equids) 
2 on the upper edge of the calcis (2 equids) 
1 on the medial side distally and on upper surface (equid) 
1 on the dorsal side close to the articulation with the astragalus ( 1 

cervid) 

Cuts are related to plantar ligament and lateral and medial liga- 
ments, with the cutting directed at dismembering the ankle joint. 

Four calcanei are chewed, three of them very heavily (Fig. 4A). 
Puncture marks are superimposed over cut-marks and percussion 
marks on one of these calcanei (M50029), which indicates that 



carnivore activity occurred after human. Chewing marks are pits on 
the surface (type a, average 2.0mm, N = 6), grooves on surface (type 
b, average 1 .6mm, N = 7) and only one of type c ( 1 .5 mm) and one of 
type h (3mm). 

Summary of calcanei. Total 8 specimens, 7 equid, 1 cervid 
Cut-marks: 5 specimens (4 equids, 1 cervids). 
Percussion: 1 specimen (1 equid) 

Astragalus: there are ten specimens of equid astragali (left 5, right 
5, MNI 5) and five cervid astragali. Nearly all specimens are com- 
plete. Cut-marks are present on five of the equid astragali and two of 
the cervid astragali: 
5 astragali have cuts on the medial condyle, medial side (3 equids and 

2 cervids) 
4 astragali have cuts on the medial surface of the body, and on the 

proximal and/or distal tuberosities (2 equids, 2 cervids) 
2 astragali have cuts on the central trochlear ridge ( 1 equid, 1 cervid) 

1 equid astragalus has cuts on the lateral condyle, medial side 

2 astragali have cuts on the lateral side of the lateral condyle ( 1 equid, 

1 cervid) 
1 equid astragalus has cuts on the posterior surface of the body. 

Cut-marks are most abundant at the medial condyle and medial 
surface of the body (Fig. 4B). The medial surface bears on its distal 
part a large tuberosity and on its proximal part a smaller one for the 



66 



P. ANDREWS AND Y. FERNANDEZ- JALVO 


















■H 






H : 












m ^ 


w 


■■ 




i 


WW ^1 


b| 


^ '.^ 


m^L. 


^J^gH| 


^ i* L J 



Fig. 5 A, Distal metapodial of Equus ferns, M50043. Deeply incised cut- 
marks can be seen around the edge of the articular surface. B, Three 
distal metapodials of Equus ferus, from left to right ventral view of 
M49834, dorsal views of M49977 & 49950. In each case, the shaft is 
split up to the articular surface with oblique fractures, curved, smooth, 
and shaft circumference 2. On M49834 there are cut-marks on the 
terminal end of the central ridge of the trochlea and a conchoidal scar on 
the dorsal side of the oblique fracture (not seen here); M49977 has 
percussion marks on the central ridge of the trochlea, visible here on the 
dorsal aspect as discolouration of the articular surface, conchoidal scars 
along the oblique break, and cut-marks on the shaft; M49950 also has 
cut-marks on the shaft along the ridge bordering the post-articular sulcus 
and conchoidal scars along the oblique break, and there are extensive 
percussion marks on the distal (terminal) part of the articular surface. 
These modifications appear to be concerned with breakage of the shaft 
for extraction of marrow and disarticulation of the foot. A, x 0.7; B, 
x0.4. 

attachment of the medial ligament of the hock joint. Cuts are 
therefore aimed at disarticulating the tarsal bones and tibia. Most of 
the marks were distributed in clusters of short incisions but no chops 
or percussion marks have been recorded. The lateral surface is 
smaller and has a wide rough fossa in which the lateral ligament is 
attached, and only isolated incisions have been found on the lateral 
trochlea. No human activity has been observed on these bones, so 
that there is no evidence of percussion or conchoidal scars. There is 
also no evidence of peeling, flakes removed or adhered flakes. 

Carnivore chewing marks are present on two specimens, one of 
them heavily chewed (M50003, lacking cut-marks). The chewing 
marks are mainly on the articulation with the calcaneus (type b, 
average 1.9mm, N = 2 and type c, average 2.8mm, N = 14) 



Summary of astragali. Total 15 specimens, 10 equid, 5 cervid 
Cut-marks: 7 specimens (5 equid, 2 cervid). 

Third tarsals: Seven specimens of equid tarsals have been seen, 
all complete. Cut-marks are present only on two specimens, trans- 
verse incisions across the dorsal surface. Three of the tarsals are 
slightly cracked by weathering, and two have manganese stains. 

Other podials: There are a magnum and a scaphoid of equid, two 
naviculars ( 1 equid, 1 cervid) and a central tarsal of horse, all of them 
complete. Human-induced damage is evident on the scaphoid that 
has cut-marks on the dorsal surface, and on the central tarsal that 
bears an adhered flake on a lateral broken surface on the articular 
dorsal ridge. All of them are slightly cracked on surface. 

Metapodials: There are five human metatarsals, twenty four 
metapodials of equids and six of cervids. They have mostly come 
from the earlier excavations, with only three of the human metapodials 
coming from the 1987 excavations. 

Human metapodials are mainly shafts, sometimes with one end. 
They have no cut-marks on their surface or articulations. Animal 
metapodials are all distal ends, except for 1 3 lateral metapodials of 
equid that are complete. All human and equid lateral metapodials 
have length category 3 (almost complete). Medial metapodials have 
circumference almost complete with the exception of three of them 
that have circumference 2, but they have length values that are 
mostly category 1 (less than l/4th of the length) or category 2 (less 
than half of the original length) (Villa and Mahieu 1991 ). Cut-marks 
are present on 1 8 of the 30 animal metapodials (6 cervids, 12equids), 
but they have been found on none of the humans. Cuts are located on 
the trochlea, all round the articulation (Fig. 5A) or on the dorsal/ 
ventral surfaces close to the articulation. Medial metapodials of 
horse have a consistent pattern of breakage (Fig. 5B) indicated by 
breakage on the shaft close to the distal articular end and extensive 
percussion marks providing similar bone fragments. 

Human metatarsals are all crushed on one or both ends. The ends 
show evidence of chewing marks, peeling or percussion marks on the 
edge of the articulation. Two human metapodials appear chewed (a 
5th and a 2nd left), although no actual clear puncture mark can be 
measured. The 2nd left human metatarsal shows strong similarities 
with a suid rib experimentally chewed by humans (Fig. 6). In 
contrast to this, no carnivore tooth marks have been recorded on any 
other animal metapodial surface. 

With regard to lateral metapodials of equids, they show similar 
crushing on articular ends to that seen in humans. Lateral horse 
metapodials have a consistent pattern with percussion marks on 
proximal ends on the outer (lateral) surfaces and only rarely on the 
inner articular surface with the medial metapodial (only 1 specimen). 
Percussion marks appear on one specimen distally as well. Some- 
times, percussions are associated with chop marks (2 cases) and/or 
cut-marks (3 cases) trans versally to the length of the bone. Percussion 
marks are located on the lateral tuberosity of the proximal end. One of 
the bones has three chewing marks, type b averaging 1.4mm, N = 3. 

The length of metapodials is variable, but none has been split 
longitudinally. Percussion marks on 21 of the metapodials, however, 
are located at one side (ventral or dorsal) or distributed on lateral, 
ventral and dorsal surfaces or, more exceptionally on the articula- 
tion. A distal metapodial of cervid has a percussion triangular shape 
as observed in mandibles (see below). Some metapodials have been 
seen to have scratches on the opposite side of percussion marks, 
probably as result of anvil effect on the bone during breakage. Four 
metapodials show conchoidal scars and flakes removed on the 
broken edge. 



CANNIBALISM IN BRITAIN: TAPHONOMY OF FAUNAL AND HUMAN REMAINS FROM GOUGH'S CAVE 



67 



p» 1 






^H 


1 / 


' ■ ^ . 


mi 










Fig. 6 A, a suid metapodial; and B, a second left human metatarsal, 
GC87-30. The suid metapodial was experimentally chewed by humans. 
The extensive fracturing of the proximal ends, with depressed flakes of 
bone and splaying of the ends, is extremely similar in both bones, and 
may indicate human chewing on the metatarsal from Gough's Cave; 
x 1.2. 

Only three metapodials have trampling marks and five have 
evidence of wet abrasion. One metapodial is weathered in stage 3 or 
4 and two are in stage 1 or 2. Manganese oxide stains affects 1 of the 
35 metapodials. 

Summary of metapodials. Total 35 specimens, 5 human, 24 equid, 6 

cervid 

Cut-marks: 18 specimens (12 equids, 6 cervids). 

Percussion: 22 specimens (1 human from the 1987 collection, 19 

equids (11 lateral metapodials), 2 cervids) 

Conchoidal scars: 4 specimens (4 equids) 

Flakes removed: 2 specimens (1 equid, 1 cervid) 

Peeling: 1 specimen (1 human) 

Phalanges: There are 22 proximal phalanges, one human, 20 horse 
and one cervid. The only human phalanx here studied is from the 
hand and it has the proximal end smashed and the distal end intact. 
This is similar to patterns observed in the human collection of 
Atapuerca, Mancos and the Anasazi pueblos (Andrews & Fernandez- 
Jalvo 1997). 

Most horse proximal phalanges are intact except for M49945, 
which has the proximal end removed by heavy percussion impacts. 
A flake has been removed on the medial edge, and the shaft is 
cracked both longitudinally and transversely with percussion marks 
on the broken edge. Another horse phalanx (M49788) is complete, 
but shows very heavy percussion marks on ventral (palmar) surface: 
two multiple marks on the distal articular surface (Fig. 7B) and two 
extensive percussion marks on the proximal articular end. In general, 
percussion marks may occur on the lateral, dorsal or ventral surfaces, 
as well as proximal and distal ends. Cut-marks are oblique to the 



length of the shaft on both dorsal and especially on ventral surfaces 
(Fig. 7 A). Carnivore chewing appear on two phalanges (M 49787) 
showing four carnivore pits on the surface (type a, average 1 .7mm, N 
= 4), grooves on the surface (type b, average 1.4mm, N = 14), and 
eight carnivore pits on articular surfaces (type c, average 1 .65mm, N 
— 8). The average width of carnivore gnawing grooves that affect the 
proximal phalanx M49958 is 0.72mm. Manganese staining affects 
eight proximal horse phalanges on the surface. There is no evidence 
of water damage observed on other phalanges from this study 
collection. The only proximal cervid phalanx is longitudinally bro- 
ken with grooves and percussion marks on both sides of the fracture. 
Cuts affect the phalanx on the dorsal surface and on the articulation. 

There are 13 middle phalanges, 1 1 of horse and two of cervid. 
Fewer cut-marks and percussion marks are present on the middle 
phalanges, and where they occur they tend to be at the end of the 
bones near the articular surfaces. Two of the horse phalanges show 
percussion marks, but both of them are complete. One of them 
(M49921) has extensive percussion all around the dorsal (proxi- 
mally on articular surface) and lateral surfaces. The horse phalanx 
labelled as number 23 from the excavations of 1987 shows also 
extensive percussion marks on the dorsal surface. Cut-marks are 
abundant and very marked on M50030 (Figs 7B, 7C) affecting dorsal 
and ventral surfaces on the shafts, proximal and distal ends and 
articular surfaces. Manganese stains cover all over the surface of 
three medial horse phalanges. Both second phalanges of cervid are 
complete. Phalanx M49758 shows deep cuts on the ventral side on 
the shaft and a couple of small incisions on the lateral side of the 
articulation (distal end). 

There are 18 terminal phalanges of horse and one of cervid. 
Eleven of the horse phalanges have percussion marks and cut-marks 
on the flexor surface, hoof surface and dorsal surface (Fig. 8). Six of 
horse and the cervid phalanges have no modifications. Cuts on the 
ventral side affect the deep flexor tendon, although this would seem 
to be an uncommon area of cutting related to dismemberment of the 
hoof. Two horse phalanges (M49959, M49879) show water damage. 

Summary of phalanges . Total 54 specimens, 1 human, 49 equid, 4 

cervid 

Cut-marks: 31 specimens (29 equid,. 2 cervid) 

Percussion marks: 29 specimens (27 equid, 2 cervid) 

Anatomical elements: axial skeleton 



Ribs: The human ribs from Gough's Cave are attributed to three 
individuals (Churchill 2000). The first individual has lightly con- 
structed ribs and must have been relatively small. The associations 
between ribs are based on size, curvature and morphology of the 
iliocostal line, which is superoinferiorly compressed in this indi- 
vidual (Churchill 2000). The heads of all but one rib are missing from 
this individual, perhaps because it represents an immature indi- 
vidual. The ribs of the second individual are more robust, with 
heavier muscle markings, and again the heads are missing from all 
but one rib. The ribs from these two individuals were found in partial 
association during the 1986-1987 excavation, in close proximity 
although not articulated (Fig. 9, data from 1987 excavation). A third 
individual is represented by six fragments forming parts of three ribs 
from the left side, and a further ten fragments that could not be 
further identified are also present (Churchill 2000). The 40 human 
ribs contrast with only five of large mammal. Individual two is the 
more complete, with 18 ribs, 6 of them complete. Individual one has 
16 ribs, 6 of them complete. Human induced damage on the ribs is 
very extensive (Table 3), with 30 of the 45 total number of ribs 
(animal and human) showing human-induced damage. Of the 



68 



P. ANDREWS AND Y. FERNANDEZ-JALVO 




Fig. 7 A, First phalanx of Equusferus M49730. Two areas of cut-marks are present, both on the ventral surface, one laterally and the other proximally. B, 
First phalanx of Equusferus M49788. Very heavy percussion marks are present on the ventral (palmar) surface of the distal articulation, seen as two 
rosette-shaped multiple marks with a single additional mark laterally, and at the proximal end there are two extensive areas of percussion damage 
ventrally, on either side of the proximal articular surface and extending round on to this surface. C, Middle phalanx of Equusferus, M50030. A series of 
distinct cut-marks cross the dorsal surface on the shaft and the medial edge of the distal articular surface. D, SEM micrographs of a set of stone tool cut- 
marks of a second phalanx of equid (M49999). These marks are related to meat filleting. The cut-marks are characterised by V-shaped sections, 
microstriations running linearly along the length of the cut, lateral and more superficial cut ( 'shoulder effect' Shipman and Rose 1 98 1 ) and irregular 
displaced bone on the side of the striations caused by resistance of the bone to the cut friction ( 'herzinian cones' Bromage and Boyde, 1984). A, x 0.9; B, 
x0.85;C,x 1.3. 



unmarked ribs, only 2 of the 15 are complete. Cut-marks are present 
on 25 of the ribs and occur on the shafts as well as both caudal and 
sternal ends. Extensive peeling is seen on the human ribs, sometimes 
on both ends of the fragment and sometimes also on the shaft at the 
inferior border or broken edges. Percussion marks are frequent on 
most of the ribs (Fig. 10B). 

There is one case of percussion damage on associated human ribs 
that suggest that this activity took place when the ribs were still 
anatomically joined together. There are two percussion/chop marks 
that coincide between the inferior border of rib 5 and the superior 
border of rib 6 of individual 2 (Fig. 10 B). In addition there is a 
massive chop mark on the superior border of rib 5 coinciding with a 
percussion mark on the inferior border of rib 4 from the same 



individual. Rib 4 also has extensive peeling all along the inferior 
border on the outer surface (Fig. 1 1 ). The inner surfaces of the ribs 
are affected by cut-marks on 5 human ribs from both individual 1 and 
2, and similar marks are seen on a large mammal rib that also bears 
percussion marks. In any case, the outer surface of the ribs is the most 
affected area by intensive cutting, percussion and peeling. 

Cut-marks on ribs are mainly oblique to the long axis of the bone, 
but also longitudinal and sometimes transverse around the head of 
the ribs on both animal (Fig. 12A) and human ribs (Fig. 12B). 
Scratches related to anvil-hammerstone effect have also been 
observed on at least two ribs. Fig. 1 3 shows the actual number of cut- 
marks found on all human ribs from Gough's Cave. 

Tooth marks are recorded on only two human ribs, but they could 



CANNIBALISM IN BRITAIN: TAPHONOMY OF FAUNAL AND HUMAN REMAINS FROM GOUGH'S CAVE 



69 



Table 3 Human induced modifications observed on ribs and clavicles. The ribs are identified to individual 1 or 2, their number and left or right. Modifica- 
tions are shown according to their position on the bones (internal surface, outer surface, caudal end, mid-haft, sternal end, and inferior or superior 
borders). Modifications are identified as c = cutmark; p = peeling; perc = percussion/chop marks. Sequences of modifications are given in order of 
importance. 



Register 
number: 
RIBS 


Identity 


Side 


Internal 
surface 


Outer 
surface 


Caudal 


Shaft 


Sternum 


Inferior 


Superior 


M54016 


human 


ind.2 


V L 




c 




c 








M54017 


human 


ind.2 


2 nd L 




p/c 




p/c 




P 


c 


M54018 


human 


ind.2 


3 rd L 




P 


P 










M54019 


human 


ind.2 


4 th 




c/p/perc 




p/perc 


c 


perc/p 


c 


M54020 


human 


ind.2 


5"'L 




perc 




perc 




perc 


perc 


M54021 


human 


ind.2 


7"'L 




(chop)/c/p 


c/perc/p 


perc(chop) 




perc 




M54022 


human 


ind.2 


6'" L (2/3 rib) 




perc/c 


c 


perc(chop) 


c 




perc(chop) 


M 54023 


human 


ind.2 


8 ,h -9' h L (head rib) 




c 


c 






c 




M 54024 


human 


ind.2 


8 lh -9 lh L (body frag) 




c 




c 








M54025 


human 


ind.2 


8' h -9 lh L (body frag) 
















M54026 


human 


ind.2 


2 nd R (1/2 body) 
















M54027 


human 


ind.2 


3 rd R 




perc 




perc 








M54028 


human 


ind.2 


4"'R 




c/p/perc 




perc 


c/p 




chop 


M54029 


human 


ind.2 


5 ,h R 




c/perc 


perc 




c 


perc 




M54030 


human 


ind.2 


6"' R (part body) 




c 




c 






c 


M54031 


human 


ind.2 


7"'-9' h R(headofrib) 


c 




c 










M54032 


human 


ind.2 


10 lh R (part body) 
















M 54001 


human 


ind.l 


1 S, L 




c 


c 






c 


c 


M54002 


human 


ind.l 


2 nd L (caudal end) 
















M 54003 


human 


ind.l 


3 rd L (caudal end) 




P 




P 




P 




M54004 


human 


ind.l 


4"'L 
















M54005 


human 


ind.l 


5'"L 




c 






c 






M54006 


human 


ind.l 


6 ll, L 
















M 54007 


human 


ind.l 


7 .h_ 9 .h L (frag b , ade) 




c/perc 




c 






perc 


M54008 


human 


ind.l 


12 lh L 




P 












M 54009 


human 


ind.l 


2 nd R 


c 




c 






c 




M54010 


human 


ind.l 


4"'R 


c 






c 








M54011 


human 


ind.l 


5 lh R 
















M54012 


human 


ind.l 


6 lh R (caudal end) 


c 


c 


c 






c 


c 


M54013 


human 


ind.l 


7 ll, -9 l " R (shaft) 
















M54014 


human 


ind.l 


11"' R 


c 


c 


c 




c 


c 


c 


M54015 


human 


ind.l 


12 ,h R (part blade) 




c/p 




c/p 




P 


c 


M 5403 3 


human 


















M54034 


human 






c 






c 


c 


c 


M54035 


human 


















M54036 


human 


















M54038 


human 


















M54040 


human 


















M54041 


human 


















M54052 


human 


2 indet. fragments 
















GC86 28 


equid-cervid 






c 






c 




c 


GC'86 28 


equid-cervid 
equid-cervid 


L 




perc/c 
perc/c 


perc 


perc/c 
c 




P 




GC'89 3 


equid-cervid 




perc/c 


c 


perc/c 






P 




GC90 184 


equid-cervid 






c 












CLAVICLES 




















M54053 


human 


L 


c 


c 


c 


c 


c 


c 


c 


M54054 


human 


R 




c/p 


P 




c 


scraping 




M54055 


human 


R 


c 


c 


c 






c 


c 



not be measured. It is possible that these chewing marks could be 
human in origin (1 human rib). Carnivore chewing has also been 
recorded on one deer rib, type a average 1.9, N = 6, and type b 
average 1 .3, N = 5. Trampling marks have been seen on 4 human ribs. 
Six ribs are weathered, but only to stage 1 . One of the human ribs is 
affected by weathering on the outer side but the inner side looks 
fresh. Manganese oxide stains are present on one large mammal rib, 
and one human rib has root-marks on its surface. 

Summary of ribs. Total 45 specimens, 40 human, 5 large mammal 



Cut-marks: 25 specimens (20 humans, 5 large mammals) 
Percussion: 1 1 specimens (8 humans, 3 large mammals) 
Peeling: 8 specimens (8 humans) 

Clavicles: There are three specimens of human clavicle, two of 
them almost complete (Fig. 14). They have cut-marks on both the 
inner and outer surfaces, caudal, shaft and sternum surfaces, and on 
both the inferior and superior edges (Table 3). There are no percus- 
sion marks, anvil-hammerstone scratches, conchoidal or flakes that 
may suggest breakage of this anatomical element, though peeling is 



70 



P. ANDREWS AND Y. FERNANDEZ-JALVO 




Fig. 8 Two views of distal phalanges of Equusferus. A, dorsal view, with 
transverse cut-marks across the body and dorsal edge of the proximal 
articulation; B, an enlarged view of these cut-marks. A, x 1 . 1 : B, x 2.2. 

present in one of the clavicles (M54054) on the caudal outer surface 
indicating bending of the bone to dismember or detach the clavicle. 
Scraping also occurs on this clavicle. 

Vertebrae: There are ten human vertebrae in the Gough's Cave 
material. Most of these were cervical vertebrae, but there were four 
fragments of thoracic vertebrae, three neural arches and part of one 
body. It is possible that nine of the vertebrae could be from a single 
individual (Churchill 2000), and since they were found with the two 
partial sets of ribs it is likely that they came from one of these 
individuals. We were able to study seven of the vertebrae ( 1 axis, 3 
cervicals and 3 thoracics), seven of equid ( 1 atlas, 1 axis, 4 cervicals 
and sacrum). There are five cervid vertebrae (3 cervicals, 1 thoracic 
and the 1st caudal). Most vertebrae are affected by human induced 
damage (Fig. 15), with only two human and two cervid vertebrae that 
are intact and undamaged. Neural arches are broken in two of the 
human vertebrae. The vertebral bodies from humans, equids and 
cervids have cut-marks or percussion marks, especially on the ante- 
rior part of the body. The human axis (M54042, Fig. 1 6A) has cuts on 
the anterior side of the body along the insertion of the stylohyoid 
muscle (Fig. 16B). Most transverse processes on human, equid, and 
cervid vertebrae are broken or cut on the posterior part of the 
vertebra. One human vertebra shows peeling on the transverse 
processes. The laminae of the vertebrae, especially those of equids, 



but also of humans and cervids, are also broken (peeled apart in 
humans) or cut on one side or all round the spinous process. 

Tooth marks are abundant on the vertebrae, with five specimens 
showing chewing or tooth marks, but only a few of these were clear 
enough to be measured on the equid sacrum. The type of tooth marks 
are pits on bone surface (a, average 2mm, N = 2), grooves on bone 
surfaces (b, average 1 .6 mm, N = 4), and tooth print (g, 2.9 + 2.9, total 
length 7.9 mm). Trampling marks have been seen on two cervid 
vertebrae and manganese oxide stains on three vertebrae of equid. 

Summary of vertebrae. Total 19 specimens, 7 human, 7 equid, 5 

cervid 

Cut-marks: 1 1 specimens (3 human, 6 equid, 2 cervid) 

Percussion: 2 specimens ( 1 human, 1 equid) 

Peeling: 2 specimens (2 human) 

Tooth marks: 5 specimens (4 equid, 1 cervid chew mark) 

Anatomical elements: flat bones 

Pelvis: Two equid pelves are the only specimens found of this 
element. One specimen has the ileum and part of the acetabulum 
preserved, the other is almost complete with the pubis recently 
broken. Cut-marks (Fig. 17) are concentrated along the spine and 
along the posterior edge on the dorsal side of the ileum, as well as on 
the ridge below the acetabulum, and chop marks are present on the 
spine near the acetabulum. The most complete pelvis has cut-marks 
on similar areas but also including the ischium on both sides (dorsal 
and ventral). Trampling occurs extensively on one of the pelves, and 
manganese oxide stains occur on both. Both pelves also have carni- 
vore damage, mainly on the ischium and ventral side of the ileum 
proximal edge. Types of tooth marks are as follows (Andrews & 
Fernandez-Jalvo 1997): a (pit marks on bone surface) average 1.5 
mm, N = 5; b (grooves on bone surface) average 1.5 mm, N = 4; c 
(punctures on anatomical borders) average 3.5 mm. N = 2. 

Summary of Pelves. Total 2 equid specimens 
Cut-marks: 2 specimens 

Scapula: The scapula is also poorly represented in the collection, 
with just four fragments from humans. Cut-marks and scraping 
marks are extensive on all four scapula fragments. The scapula 
M54057 which was partially recovered in 1927 has been completed 
by refitting fragments recovered in 1 987. The cuts are mixed on most 
scapulae with trampling marks (Fig. 18A). Some incisions are 
definitively human made because they bend without interruptions 
and pass over and around curvatures in the bone. They are present on 
both dorsal and ventral sides. Cuts also occur over areas protected by 
the curvature of the bone (eg. between acromium and the scapula 
neck: Fig. 18B). In these positions, trampling is impossible because 
they are deeply recessed, and further the marks are deeply incised, 
which is not usual in trampling marks (Andrews & Cook 1 985). Cuts 
show a random distribution by direction related to the strong muscle 
attachments. Scrapes occur on the deeply concave angle between the 
spine and the infraspinatous fossa. Peeling occurs laterally on two 
specimens (M54057 and M54059) and the latter also has percussion 
marks on the spine, probably as result of dismembering processes of 
the humerus from the scapula. Only one scapula (M54057) shows 
evidence of weathering which is located along the infraglenoid 
tubercles along the lateral margin, indicating the scapula was resting 
with the spine down in the soil. 

Summary of scapulae. Total 4 specimens all human 
Cut-marks: 4 specimens 
Percussion: 1 specimen 
Peeling: 2 specimens 



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71 




Fig. 9 The Gough's Cave excavation showing some of the human ribs in individual 2 being excavated beneath the overhang of the side wall. 



Anatomical elements: Cranial 

Mandibles: There are three adult human mandibles in the collec- 
tion, counted as six hemi-mandibles, 1 2 specimens of equids. and 9 of 
cervid. All the mandibles are heavily damaged. The equid mandibles 
consist either of symphysis (50%) or alveolar fragments (50%) with 
no inferior border or ascending ramus. Most mandibles have some of 
the premolars and molars in situ. Cervid mandibles show a similarly 
highly destructive pattern, with broken fragments consisting only of 
portions of alveolus with no inferior border or ascending ramus. The 
only exception to this pattern is shown in Fig. 19, where a cervid 
mandible is shown with a dental series and diastema complete to the 
symphysis. Most mandibles of cervid (56%), but also some horses 
(33%), show percussion marks on the lingual side, some of them with 
massive damage (Fig. 1 9B ). Also common are cut-marks on the buccal 
side (Fig. 19A). Such strength applied to the mandible has produced 
severe damage, not only to the mandible, but also to most of the teeth, 
which are seriously crushed (Fig. 1 9B ). It is further peculiar that three 
of the cervid mandibles and one of the horse mandibles have a 
triangular shaped percussion mark (Fig. 19B), also seen on a distal 
metapodial of cervid (M49832). The triangular percussion mark is 
quite deep, suggesting a forceful stroke. 

Human mandibles are also heavily damaged in a similar way, 
although they have no percussion marks. A complete human mandi- 
ble (M54137b) has both ascending rami broken. The right 
hemi-mandible has extensive peeling on the ascending ramus, possi- 
bly related to breakage of the articular condyle, and slight breakage 
is seen along the inferior border close to the mandibular angle (lower 



angle of the mandible). The left hemi-mandible has the ascending 
ramus broken in the region of the temporal muscle insertion, (coro- 
noid process) and the articular condyle. Another hemi-mandible 
shows deep incisions at the ascending ramus (Fig. 20A) probably 
inflicted as a result of masseter muscle removal. 

A third human mandible (M54130a) shows extensive cuts on the 
lingual surface of the body (along the linea mylohyoidea) and also on 
the buccal side, and there are percussion marks on the buccal side as 
well. The left ascending ramus is broken at the articular condyle and 
the right ascending ramus is missing. The inferior border on both 
sides of the mandible is broken, and peeling is evident on the left side 
along the fracture edge. Cuts are also present on the symphysis on the 
lingual side (fossa digastrica and spina mentalis) (Fig. 20B). It is 
remarkable that one horse symphysis also shows cuts on the inner 
margin of the symphysis (Fig. 20C) similar to that seen on humans. 
This evidence is reinforced by the recovery of the hyoid of a horse 
that has percussion marks and is extensively cut. 

Trampling is evident on equid and cervid mandibles. Weathering 
has been detected on three mandibles (two of horses and one of deer) 
at stage 1 and 2/3 on one of the horses. Manganese covers four of the 
deer and horse mandibles, with the three deer mandibles are heavily 
stained. 

Summary of hemi-mandibles. Total 27 specimens: 6 human, 12 

equid, 9 cervid 

Cut-marks: 6 specimens (4 human, 3 equid, 2 cervid) 

Percussion: 13 specimens (9 equid, 4 cervid) 

Adhered flakes: 1 specimen (1 cervid) 

Peeling: 5 specimens (1 human, 3equid, 1 cervid) 



72 



P. ANDREWS AND Y. FERNANDEZ-JALVO 




Fig. 10 A, Three human ribs from individual 2, numbering from the top 
ribs 4, 5 & 6. The inferior border of rib 4 (M54019) has several 
percussion/chop marks, notably one about one third the way from the 
caudal end; the superior border of rib 5 (M54020) has a chop mark 
opposite this mark on rib 4, and it has two percussion/chop marks on its 
inferior border; the superior border of rib 6 (M554022) has a chop mark 
opposite the first and a percussion pit opposite the second. These are 
shown in context in Fig. 1 1. B, Rib of Cervus/Equus (GC86-28) with 
head intact and much of the shaft, with extensive cut-marks along the 
superior border and percussion marks on the inferior border. A, x 0.7; B, 
x0.55. 



Fig. 11 Drawing of left rib cage with the percussion and chop marks of 
ribs 4, 5 & 6 shown in the context of the whole rib cage. 




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73 




Fig. 12 A, Caudal end of Rangifer tarandus rib, no number, with cut- 
marks running transversely across the neck of the rib close to the 
articular facet of the tubercle. B, Caudal end of a human rib, M54009, 
with both ends broken and cut-marks running transversely across the 
neck of the rib and into the articular facet of the tubercle. A, x 1 .3; B, x 
1.4, 





Fig. 13 Generalized view of human rib showing the numbers of cut- 
marks found on all human ribs from Gough's Cave, and their distribution 
on the rib. 



Fig. 15 A, Axis vertebra M54042. Numerous cut-marks are present on 
this bone, particularly along the anterior surface of the body of the 
vertebra (i.e. on the front of the body, not the back). The conformation of 
these cut-marks is shown in Fig. 16. There are also cut-marks superiorly 
next to the articular surface connecting to the atlas vertebra, and in this 
region there is also peeling of the bone on the lateral aspects on both 
sides of the articulations. B, Cervical vertebra of Equus ferns (M50068) 
with cut-marks along lateral-ventral surface. Cut-marks are short and 
concentrated along the anatomical edge probably related to the 
detachment between vertebrae. A, x 1.2; B, x 0.65. 

Maxillae: There are two human palates, and three half-maxillae of 
equid and four of cervid. Two human half-maxillae, with both 
zygomatics broken, are from the same individual (M54130b), and 
they fit with mandible M54130a described above (Fig. 21 A). The 
muscle insertions of the nose and lips (levator muscles and the 
zygomatic and masseter muscles) are affected by cut-marks on the 
human specimen, which also has cut-marks on the front of the palate. 
Apart from cuts, the nasomaxillary bone is heavily modified around 
the lips and nose, and this is similar to cut-marks seen on the buccal 
surfaces of the molars (Figs 22, 23) as it is observed in other 
cannibalistic sites (Atapuerca in Spain, Fontbregoua in France and 




Fig. 14 Right clavicle M54055 diaphysis. Cut-marks are present in two places, two marks on the deltoid insertion and two also where the cortaclavicular 
ligament attaches to the shaft of the clavicle; x 1. 



74 




P. ANDREWS AND Y. FERNANDEZ-JALVO 




Fig. 16 A, Human axis vertebra M54042 showing the conformation of 
the cut-marks along the anterior surface. This is the area of attachment 
of the anterior longitudinal ligament, and in addition the superior marks 
are probably related to the detachment of the axis from the atlas, and the 
posterior marks to the detachment of the axis from the third cervical 
vertebra; x 1.2. B, Schematic drawing of human skull and upper 
vertebrae showing the disposition of two muscles that insert on the 
internal surface of the mandibular symphysis (digastric muscle) and the 
back of the skull (stylohyoid muscle). 

Anasazi pueblos in States). Breakage of the zygomatic arches is 
necessary in order to remove the temporalis muscle so as to open the 
vault for access to the brain tissues. 

Other animal maxillae are heavily broken, with only alveolar 
fragments and few premolar and molar preserved in situ. Horses 
show percussion marks on buccal and lingual sides, or only on the 
buccal. Cut-marks have not been seen on the bone of horse maxillae, 
but there are two specimens that show oblique cuts on the buccal side 
of the premolar/molar toothrow (Figs 2 IB, 22). Extensive percus- 
sion marks appear on both the lingual and the buccal sides of horse 
maxillae, and one specimen has adhered flakes also on both sides. It 
is interesting to note that one horse maxilla has peeling on the palate. 




Fig. 17 Innominate of Equusferus M50028. This bone has been 
extensively modified, with loss of the extremities and many trampling 
marks on both surfaces. There are many carnivore tooth marks on the 
upper border of the ilium, and four chop marks are also present crossing 
the spine near the superior border of the ilium. More inferiorly there are 
numerous cut-mark incisions concentrated along the spine; x 0.8. 

With regard to cervids there are cuts only on the buccal side of the 
maxilla, along the dental series (Fig. 21C). Percussion marks on the 
lingual side have only been seen on one cervid specimen. No teeth 
have cut-marks on cervid maxillae, but the teeth are heavily crushed 
on the lingual side or broken. 

Summary ofhemi-maxillae: Total 9 specimens, 2 human, 3 equid, 4 

cervid 

Cut-marks: 6 specimens (2 human, 2 equid, 2 cervid) 

Percussion: 6 specimens (2 human, 3 equid, 1 cervid) 

Adhered flakes: 1 specimen ( 1 equid) 

Peeling: 1 specimen ( 1 equid) 

Skull: There are three human calottes (frontal, parietal and most 
part of the occipital) of an adult, adolescent and child. The adult skull 
is almost complete, although the face is missing (broken at the orbital 
region and maxilla), and a frontal fragment. There are two cervid 
skulls, both broken but there is no conclusive evidence of human 
damage. One of the cervid skulls shows weathering at a stage 1. 

The human calottes show similar patterns to each other regarding 
cut-marks and percussion marks. Extensive and long cut-marks are 
present on the temporal insertions of the parietal bones on both sides 
(Figs 24A, C). Many cuts are also present on the frontal bones (Fig. 
25) and on the supraorbital ridges (insertions of the orbicular and 
superciliary muscles), as well as in the eye sockets to extract the eyes 
(Fig. 24B). On the occipital bones, cuts are also seen along the 



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75 




Fig. 18 A, Left scapula with the spine and part of the inferior blade of 
M54059. There are many trampling marks mixed in with cut-marks, and it 
is sometimes hard to distinguish them. Numerous scrapes occur in the 
deeply concave angle between the spine and the infraspinous fossa, with 
percussion marks on the edge of the inferior border. There are also 
percussion marks and peeling on the acromion process. The scrape marks 
are not straight, bending without interruption over and around curves in 
the bone surface, and they cut across several incisions perpendicular to the 
general trend of the scrapes, the cut-marks clearly preceding the scrapes. 
In the angle between the acromion and the scapular neck, cranial 
orientation, there are two oblique and deep incisions not visible on this 
view but similar to the ones in M54056. B, Right scapula M54056. Two 
incisions, one deep and the other shallow, are present in the angle between 
the acromion and the neck of the scapula, and in addition several cut- 
marks are present in the inferior angle of the scapular blace at the insertion 
of teres major. A, x 0.9; B, x 0.65. 



sutures with the parietal bones, covering the insertion area of the 
trapezius and sternocleidomastoid muscles (Figs 24D, 25B). Per- 
cussion marks appear superimposed on cut-marks on the parietal 
bones at both sides (Fig. 24C) and cuts appear interrupted by the 
broken edges. Incipient peeling has been seen on the zygomatic 
arches, and some removed flakes on the broken edges (i.e. left side of 
the sphenoid bone and occipital bones). 

Summary of skulls. Total 5 specimens = 3 human and 2 cervid) 
Cut-marks: 2 specimens (2 human) 
Percussion: 2 specimens (2 human) 
Removed flakes: 2 specimens (2 humans) 
Peeling: 2 specimens (2 human) 



DISCUSSION 

Large mammal species diversity recorded at Gough's cave is quite 
poor, with most remains assigned to three species of large mammals, 
Homo sapiens, Equus ferus and Rangifer tarandus. 

The human skeletal element proportions are generally higher than 
those of any of the other large mammals. Ribs and cranial remains 
are the best represented for humans, but paradoxically, vertebrae are 
rare, whereas for the animal bones ribs are almost absent but verte- 
brae and especially phalanges and metapodials are much better 
rbepresented. Phalanges in particular are not commonly represented 
in human occupation sites. The relative abundances of anatomical 
elements suggest a type of selection for large mammal skeletons for 
heads and distal limbs, in contrast to human skeletons, for which 
elements from the thorax (except vertebrae), heads and arms have 
been selected. 

The cranial skeleton is the most extensively damaged anatomical 
element, both in humans and large mammals (equids and cervids). 
Human skulls and faces at Gough's Cave have a higher intensity of 
cut-marks than are present on non-human animals, but contrasting 
with this, two of the human skulls are almost complete. Skull 
completeness differs greatly from site to site where cannibalism has 
been considered to be nutritional (i.e. TD6- Aurora Stratum, Spain), 
the Neandertal site of Moula-Guercy (France Defleur, etal, 1999), 
or modern ones where fire is involved such as Native American sites 




|§^>«r 






B| 


1^^^^^ — **^r^^ 


J 



Fig. 19 A, lingual, and B, buccal views of deer mandible {Rangifer tarandus), M4982 1 . The buccal view shows numerous cut-marks along the diastema, 
and the lingual view shows one and possibly two percussion marks near the alveolar border; both x 0.5. 



76 



P. ANDREWS AND Y. FERNANDEZ-JALVO 






H 1 • 

^1 




MJJM 


c| 






Fig. 20 A, Human mandible (GC'87) showing deep incisions on the ascending ramus as a result of dismemberment of the mandible probably inflicted as a 
result of masseter muscle removal. B, Detail of cut-marks on human mandible M54 1 30a. The cuts are on the mandibular symphysis on the lingual side 
along the internal ridge, at the insertion of the digastric muscle. C, Mandible of Equus ferus M49848. Cut-marks are on the mandibular symphysis on the 
lingual side, and on the lingual border of the mandibular body close to the alveolar margin and on the lingual border of the diastema. A, x 2; B, x 3; C, x 
1.2. 



(USA, Turner and Turner, 1999; White, 1992), Navatu (Fiji Islands, 
Degusta, 1999). At all these sites, damage to the human skulls is great 
and it is interpreted as the result of gaining access to the brain. The 
only other cannibalistic site where human skulls are relatively com- 
plete is at Fontbregoua (French Neolithic) where Villa et al. 
( 1986a&b) interpreted it as an element of skull ritual treatment. We 
agree with this interpretation in relation to the Gough's Cave material 
because completeness of skulls, even where they have been damaged 
by percussion and intensive cutting, is an exception to the general 
pattern of the Gough's Cave assemblage. Animal skulls are notable 
for their absence, and most other skeletal elements, with the excep- 
tion of the limb extremities and some of the human ribs, are all 
broken. The human skulls stand out as the most intact groups of 
bones that by their nature are relatively easily broken by post- 
depositional processes. 

The jaws in particular have been heavily broken and cut. Most 
large mammal jaws recovered from the site consist of alveolar 
fragments, with or without teeth. Teeth are damaged by crushing, 
especially in cervids, but also in humans. There is a peculiar butch- 



ering technique observed on these specimens consisting of intensive 
cutting on the buccal side, and strong percussion marks on the 
lingual side. These suggest dismemberment of the mandible and 
cutting of the muscles of mastication and the lip depressors. Cut- 
marks lingually, particularly on the symphysis in the digastric area 
on both equids and humans, indicate removal of the tongue. Cut- 
marks have also been found on the enamel of horse upper molars on 
the buccal side. These cuts have also been observed at other sites 
such as Abric Romani (-40,000 BR Barcelona, Spain), and here they 
have been interpreted as cutting of facial muscle attachments to 
extract the cheek. 

Damage is also seen on human jaws with breakage of zygomatic 
arches on the upper jaws, and inferior borders and ascending ramii of 
mandibles. Similar destruction of human remains also appears at 
Native American sites (White, 1992; Turner and Turner, 1999), 
Fontbregoua (Villa et al, 1986 a, b) and especially at TD6-Aurora 
Stratum (Fernandez- Jalvo et al., 1999), where no complete cranial 
element (skull vault, mandible or maxilla) has yet been found. Some 
authors have considered that this degree of intensive damage of faces 



CANNIBALISM IN BRITAIN: TAPHONOMY OF FAUNAL AND HUMAN REMAINS FROM GOUGH'S CAVE 



77 




Fig. 21 A, Human maxilla (M54130a ) and mandible (M54130a) from a 
young individual. The jaws are heavily damaged by percussion and 
cutting. The maxilla has both zygomatic arches broken and extensive 
cuts on the masseter insertion, as well as on the face above the canines 
where the lips attach, and on the palate. Cuts on the mandible are present 
on both lingual and buccal sides of the mandible. Also in the area of the 
medial pterygoid insertion and coronoid process the inferior border is 
broken and the ascending ramus broken. B, Maxilla of horse Equus 
ferus, GC89-061. The body of the maxilla is brok